1989 European Neogene Mammal Chronology

European Neogene
Mammal Chronology
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Series A: Life Sciences

European Neogene
Mammal Chronology
Edited by
Everett H. Lindsay
University of Arizona
Tucson, Arizona
Volker Fahlbusch
University of Munich
Munich, Federal Republic of Germany
and
Pierre Mein
Claude Bernard University
Lyon, France
Springer Science+ Business Media, LLC

Proceedings of a NATO Advanced Research Workshop on
European Neogene Mammal Chronology,
held May 16-20, 1988,
in Schloss Reisensburg (near GOnzburg), Federal Republic of Germany
Llbrary of Congrass Cataloglng-ln-Publlcat ton Data
European Naogane •ammal chronology 1 edited by Everett H. Lindsay,

Volkar Fahlbusch, and Piarre Mein.
p. c•. -- !NATO ASI series. Series A, Life sciences ; v.
180)
"Proceedings of a NATO Advanced Research Workshop an European
Neogene Mam1al Chronology, held May 16-20, 1988 in Schloss
Reisensburg"--T.p. versa.
"Published in cooperation with NATO Sc1ent1fic Affairs Division."
Includes bibliographical references.
ISBN 978-1-4899-2515-2 ISBN 978-1-4899-2513-8 (eBook)
DOI 10.1007/978-1-4899-2513-8
1. Mammals, Foss11--Europe--Congre sses. 2. Paleontology,
Stratigraphic--Congres ses. 3. Paleontology--Neogene- -Congresses.
4. Paleontology--Europe--C ongresse&. 5. Geology, Stratigraphic-
-Neogene--Congresses. 6. Geology--Europe--Congr esses. I. Lindsay,
Everett H. II. Fahlbusch, Volker. III. Mein, Pierre. IV. NATO
Advanced Research workshop an European Neogene Ma•mal Chronology
!1988: Schloss Reisensburg> V. North Atlantic Treaty Association.
Scientific Affairs Oivision. VI. Series.
QE881.E82 1989
569'.094--dc20 89-26648
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Originally published by Plenum Press, New York in 1989
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PREFACE
During the last ZO years great progress has been achieved in our understanding of
both earth history and vertebrate evolution. The result is that climatic/tectonic
events in earth history can now be placed in a more precise and global time frame,
that permit their evaluation as abiotic causal factors which might trigger extinction
and dispersal events in vertebrate history. Great strides have also been made in
genetics and cell biology, providing new insight into phylogenetic relationships among
many vertebrates. These new data, along with data on chronologie resolution of earth
history, provide tests of previous interpretations regarding ancestral-descendant
relationships based solely on the fossil record.
It is fitting and proper that a volume on European Neogene mammal chronology

is produced at this time, to ensure that new interpretations of vertebrate evolution
and chronology are based on the most accurate and current data. Vertebrate paleon-
tologists believe that the fossil record is the only secure data for measuring the actual
course and tempo of vertebrate evolution. Knowledge of the fossil record must keep
pace with advances in other areas of science so that inferences on vertebrate evolu-
tion are accurate and meaningful.
The rich record of fossil mammals in Europe has contributed substantially to

studies of vertebrate evolution. Notable early contributions to Cenozoic geochronology
and vertebrate evolution were made by Cuvier, Geoffroy St.-Hilaire, Darwin, Huxley,
Agassiz and others. Many of the concepts that we now use in Cenozoic geochronology
and vertebrate evolution were developed in Europe during the last century.
European mammal chronology has grown from both broad synthetic studies (e.g.,
Zittel, Thenius, Kretzoi and Kurten) plus centrally focused studies (e.g., Crusafont,
Thaler, Tobien and Azzaroli). The latter of these foundation builders are included
among the contributors to this volume. We co-editors dedicate this volume to those
vertebrate paleontologists who have helped to create the foundation of European
Cenozoic mammal chronology that we have inherited.
An international workshop was held at Schloss Reisensburg in Germany on May

16-ZO, 1988, to stimulate further knowledge of European fossil mammals. This
meeting, sponsored by NATO as an Advanced Research Workshop, had the title
"European Neogene Mammal Chronology." It was attended by 47 researchers (listed at
the end of the volume) from 15 nations, including 13 European nations. Presentations
were given in English, even though English is the native tongue for only about 1Z
percent of the participants. The topics addressed at the meeting were varied, includ-
ing principles of stratigraphy, presentation of new fossil discoveries, syntheses of
climatic and biogeographic data, update of systematics for selected groups of
European mammals, and chronology of mammal faunas outside of Europe. Discussions
were friendly and lively; conflicting perspectives were addressed and discussed openly.
Contributions to this volume reflect the diversity of topics and viewpoints

presented at the Workshop. Seven chapters (e.g., 1, 6, 7, 18, 31, 35 and 37) were
initiated after the Workshop and most of the manuscripts were revised to some degree
v
after the Workshop. Note that opm1ons regarding importance and significance in
mammalian chronology are diverse; we have attempted to include all these viewpoints
in this volume, and will leave for future historians to decide which of these viewpoints
contributed the most important or significant methods and studies for advancing
mammal chronology, especially in Europe. We will claim that this volume reflects
current state-of-the-art in knowledge of European mammal chronology.
The co-editors are indebted to the North Atlantic Treaty Organization and the
National Science Foundation for supporting this Workshop. We thank the Inter-
nationales Institut fur wissenschaftliche Zusammenarbeit e.V. Schloss Reisensburg for
providing comfortable accommodations, excellent service and tasty meals during our
Workshop. A visit to the meteor crater and fossil deposits near Steinheim was led by
E.J.P. Heizmann, to whom we are grateful. An editorial board consisting of Andrews,
Fejfar, Ginsburg, Kowalski, Qiu and the co-editors critiqued the manuscripts following
the Workshop. We were assisted in the preliminary organization of the Workshop by
Mrs. Helga Fuchs, who was unable to join us during the Workshop. Ms. Gertrud
Roessner capably assisted during and after the Workshop. We are especially grateful
to Ms. JoAnn Overs for expert typing and editorial assistance in the preparation of
camera-ready manuscripts. Mr. Jim Abbott provided expert advice and skill in design
and correction of illustrations, for which we are grateful. The co-editors express their
sincere thanks to all those mentioned above, for helping to present a stimulating
Workshop and a handsome volume.
E. H. Lindsay
V. Fahlbusch
P. Mein
vi
CONTENTS
ASPECTS OF EUROPEAN MAMMAL CHRONOLOGY
The Setting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . 1
E. Lindsay
European Neogene Marine/Continental Chronologie Correlations • • • • • • • • • • • • • 15

F. Steininger, R.L. Bernor, and V. Fahlbusch
A Biochronologic Subdivision of the European Paleogene Based on

Mammals - Report on Results of the Paleogene Symposium
held in Mainz in February 1987 • . • . . . • • • . . • . • • • . • . • . • • • . • • . • . . • • • • • 47
N. Schmidt-Kittler
The Ramblian and Aragonian: Limits, Subdivision, Geographical

and Temporal Extension • • • . • . . • • . . • • • . . • . . • • • • . • • . • • • • . • • • • • . • • • • 51
R. Daams and M. Freudenthal
New Neogene Rodent Assemblages from Anatolia (Turkey) 61

M. Siimengen, E. Unay, G. Sara~, H. de Bruijn,
I. Terlemez, and M. Giirbiiz
Updating of MN Zones • • . • • • . • . . . . • • • • • • • • • • • • • • • . • • • • • • • . • • . • • • • . • • • . • • 73
P. Mein
Muroid Rodent Biochronology of the Neogene and Quaternary • • • • • • • • • • • • • • • • 91

in Europe
0. Fejfar and W. Heinrich
Biozones or Mammal Units? Methods and Limits in Biochronology • • • • • • • • • • • • 119

c. Guerin
Large Mammal Dispersal Events at the Beginning of the
Late Villafranchian . • • . . • • • • • • • • . . • . • • . • . . • • • . • • . • • • • • • . • • • • . . . • . 131
F. Masini and D. Torre
REGIONAL PAPERS
Synthesis on the "Aquitanian" Lagomorph and Rodent Faunas of the

Aquitaine Basin (France) • • • • • • . • • • • • . • • . • • • • • • • . • . • • • • . • • . . • • • • • • . 139
M. Hugueney and M. Ringeade
The Faunas and Stratigraphical Subdivisions of the Orleanian

in the Loire Basin (France) . . . . . . • . . . . . . . . . . . . . . . . . . . . . . • . . . . . . • . • • 157
L. Ginsburg
vii
A Preliminary Mammal Zonation of the Upper Marine Molasse
of Switzer land . . . . • . . . . . . . . . . . . . . . . . . . • . . . . • . . . . . . . . . . . . . . . . . . . . 177
B. Engesser
The Faunal Succession in the Bavarian Molasse Reconsidered-

Correlation of the MN5 and MN6 Faunas • • • • • • • • • • • • • • • • • • • • • • • • • • • • 181
K. Heissig
Stratigraphy of Neogene Mammals of Poland 193

K. Kowalski
The Neogene VP Sites of Czechoslovakia: A Contribution

to the Neogene Terrestric Biostratigraphy of Europe
Based on Rodents • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 211
0. Fejfar
FAUNAL DATUM PAPERS
The "Proboscidean Datum Event:" How Many Proboscideans

and How Many Events? . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . • . . . Z37
P. Tassy
The Proboscideans Data, Age, and Paleogeography: Evidence

from the Miocene of Lisbon • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 253
M. Antunes
Patterns of Old World Hipparionine Evolutionary Diversification

and Biogeographic Extension • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 263
R.L. Bernor, H. Tobien, and M. Woodburne
The Hipparions of the Lower Axios Valley {Macedonia, Greece).

Implications for the Neogene Stratigraphy and the
Evolution of Hipparions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 321
G. Koufos
The Genus Eguus in Europe • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 33 9

A. Azzaroli
BIOGEOGRAPHIC SYNTHESIS
Bioevents and Mammal Successions in the Spanish Miocene • • • • • • • • • • • • • • • • • • 357

S. Moya-Sola and J. Agustf
The Miocene Rodent Succession in Eastern Spain: A

Zoogeographical Appraisal • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 375
J. Agustf
Gundersheim-Findling, a Ruscinian Rodent Fauna of Asian

Affinities from Germany • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 405
G. Storch and 0. Fejfar
Dynamics of Old World Biogeographic Realms during the

Neogene: Implications for Biostratigraphy • • • • • • • • • • • • • • • • • • • • • • • • • • 413
M. Pickford
PALEOECOLOGICAL SYNTHESIS
Miocene Paleoecology of Pasalar, Turkey • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 443

B. Alpagut, P. Andrews, and L. Martin
viii
Taphonomic and Sedimentary Factors in the Fossil Record
of Mammals • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 461
M.A. Alvarez Sierra, M. Dfaz Molina, J .I. Lacomb a,
and N. L6pez Martmez
Relations Between Paleoclimatology and Plio-Pleistocene

Biostratigraphic Data in West European Countries , , , • , • , , •••• , , , •••• , 475
M. Bonifay
Small Mammal Taphonomy 487

P. Andrews
MAGNETOSTRATIGRAPHIC APPLICATIONS
Hipparion Datum and its Chronologie Evidence in the

Mediterranean Area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 495
S. Sen
The Magnetic Stratigraphy of the Late Miocene Sediments

of the Gabriel Basin, Spain ••••••• , , • , •• , , , • , , •• , ••• , , , •••• , •••••• , 507
N. Opdyke, P. Mein, E. Moissenet, A. P~ez-Gonz~ez,
E. Lindsay, and M. Petko
Preliminary Magnetostratigraphic Results of Some Neogene

Mammal Localities from Anatolia (Turkey).••• , , ••• , , •••• , , • , • , , , • , , , 515
C.G. Langereis, S. Sen, M. Siimengen, and E. Unay
SEQUENCES OUTSIDE EUROPE
The Chinese Neogene Mammalian Biochronology- Its Correlation

with the European Neogene Mammalian Zonation •• , •• , ••••• , , •••• , • • 527
Z. Qiu
Key Biostratigraphic Events in the Siwalik Sequence •• , •• , ••• , , , •• , • , • • • • • • • 557

J. Barry and L. Flynn
Quo Vadis, Antemus? The Siwalik Muroid Record 573

L. Jacobs, L. Flynn, W. Downs, and J. Barry
The African Dimension in European Early Miocene Mammal Faunas ••• , • • • • • • • 587
R. Savage
Development and Application of Land Mammal Ages in

North America and Europe, A Comparison • , , •••• , ••••• , ••••••••• , , , 601
E. Lindsay and R. Tedford
NEW PERSPECTIVES
The Past, the Present, and the Future 625

V. Fahlbusch and P. Mein
Contributors . . . . . . . . . • . . • . . . . . . . . . . . • . . . . . . . . • • . . . . . . . . . . • . . . . . • . • . . • . 6Z9
Subject Index • • . . . . . . . . . . . . . • . . . . . • . . . . . . . . . . . . . . . . . . • • . . • . . . . . . . . . . . • 631
Taxonomic Index (Mammal Genera) . •• . . . •. . . • . . •. •. . . •. . . . . • . • . . . • . . •. . . 651
ix
THESETI'ING
Everett H. Lindsay
Department of Geosciences
Tucson, Arizona 85721, U.S.A.
INTRODUCTION
Vertebrate paleontology represents a blend of the biologic and geologic

sciences. This blend is often reflected in individual works, with some works showing
greater emphasis and understanding of systematics and evolution; other works showing
greater interest and concern for biostratigraphy, biogeography, and taphonomy. A
result of this blend is that vertebrate paleontology has always been a very stimulating
and creative discipline, with new ideas and techniques rapidly replacing older
concepts. Another result is the absence of a singular, recognized curriculum for the
study of vertebrate paleontology. Much of the training of many vertebrate paleontol-
ogists is strongly biased by his or her parent scientific discipline, or much of it is
informal. One further result is that precise communication among and between verte-
brate paleontologists has been hindered by poorly understood and loosely defined terms
and concepts. These latter characteristics are displayed most openly when vertebrate
paleontologists f'rom separate continents come together to discuss their works.
The communication problem is not unique to vertebrate paleontologists, nor is it

an especially critical problem. However, it must be corrected or it will become a
critical problem. I am probably more aware of the communication problem than many
of my European colleagues, and will take this opportunity to identify the problem by
discussing some confusing multiple applications of terms and concepts used by verte-
brate paleontologists.
In this contribution I review some of the basic strategies used by vertebrate

paleontologists in dating a fossil assemblage, then discuss several terms and concepts
frequently used and misused by vertebrate paleontologists. I conclude this chapter
with an overview of mammal chronology from a global perspective. Throughout the
chapter I will attempt to identify where other contributions in this volume relate to
the concepts and terms discussed here. This exercise has two purposes: (a) to identify
the communication problem mentioned above, and (b) to point out the origin of some
of the confusing multiple applications of terms and concepts. It is not my purpose to
correct the terminology used by my colleagues, although I must admit guilt to making
numerous capricious grammatical suggestions on too many contributions to this
volume. Those who know me are aware that I have made more than my share of
errors. So, let me emphasize that my bias comes from experience rather than from
principle.
European Neogene Mammal Chronology

Edited by E. H. Lindsay eta/.
Plenum Press, New York, 1990
THE BASIC STRATEGIES
One of the most interesting attributes of any fossil is its age. Indeed, the
question most frequently asked about a fossil is - "How old is it?" Paleontologists use
numerous sources of information, some from the actual fossil and others from the
deposit that yielded the fossil, to answer that question. The three basic strategies
used by vertebrate paleontologists to establish the age of a fossil assemblage are:
(1) stratigraphic superposition, (Z) stage of evolution, and (3) mammal dispersal
events. Vertebrate paleontologists are fortunate in having several strategies rather
than one; in most instances, several lines of evidence operate simultaneously. When
the data from one strategy complement and corroborate another, the confidence level
for age determination increases dramatically. Figure 1 shows a conceptual relation-
ship between these strategies. As seen in figure 1, the targeted age can be reached
via superposition, stage of evolution, or dispersal events but the determination will
more likely hit the mark if constrained by several of these strategies. Superposition is
tied to biostratigraphy and is most useful if supported by magnetostratigraphy and/or
isotopic dating. Stage of evolution is tied to biochronology and depends on well
defined fossil lineages, preferably several lineages. Dispersal events depend on knowl-
edge of faunas outside the area under study; hence they relate to geochronology and
major geologic phenomena, such as globa~ sea level changes and/or climatic change.
Stratigraphic Superposition
Stratigraphic superposition is the fundamental strategy upon which the Geologic

Time Scale was established and subsequently testt'ld. Superposition is most applicable
where thick, relatively continuous, and well exposed sedimentary rocks occur. These
features are best represented in marine rock sequences; they also occur in terrestrial
rock sequences, but never as frequently nor as well developed as in marine
sequences. Superposition is invariably utilized wherever it occurs in terrestrial
sections, and frequently these superposed sections become reference sections for
shorter, less complete deposits of similar age. The development of land mammal ages
in western North America was enhanced because terrestrial deposits that yielded
diagnostic fossils were occasionally superposed. Some superposed strata in North
America became primary reference sections (e.g., the Paleocene Nacimiento Fm. and
the Eocene Bridger Fm.) for two basic reasons: They have yielded abundant diagnostic
Fig. 1. Three ways of approaching the

age of a fossil mammal assem-
blage.
2
fossils, and the fossils can be placed in an ordered stratigraphic sequence. Superposi-
tion is now demonstrable for all the North American land mammal ages, but that
potential had merely been inferred or suggested when the North American land
mammal ages were proposed less than fifty years ago.
Superposition is virtually a requirement for all biostratigraphic reference

systems; it is only a desired characteristic for biochronologic systems, such as the
North American land mammal ages. Most of the Neogene mammal faunas of Europe
lack a stratigraphic context where superposition of faunas can be demonstrated. There
are exceptions to this generalized statement, as in some basins of Spain and Greece
where superposition of mammal faunas are now well established. The contributions to
this volume by Daams and Freudenthal (Chapter 4) and by Sumengen et al. (Chapter 5)
elaborate the current "state of the art" for biostratigraphic resolution in some fossil-
rich basins of Spain and Turkey, respectively. The superposed mammal faunas from
the areas mentioned above may serve as references for European Neogene mammal
chronology; however, the location of these superposed faunas is peripheral rather than
central to the majority of European Neogene mammal faunas, and are likely to be
slightly diachronic relative to similar faunas in central Europe. This problem is
temporary, however, if the peripheral faunas can eventually be placed in a paleomag-
netic framework wherein they will rigorously test for synchrony.
Stage of Evolution
Stage of evolution, the second basic strategy for assigning an age to mammal
faunas, is a very poorly defined and poorly understood strategy. The concept was
discussed briefly at the 1988 Reisensburg Workshop and several contributions, espe-
cially Mein (Chapter 6) and Fejfar and Heinrich (Chapter 7), provide better character-
ization and specific examples of this important method. Stage of evolution has always
been well known and widely applied by vertebrate paleontologists, but it has frequent-
ly been misunderstood and/or attacked by neontologists, invertebrate paleontologists,
and others, without evoking a response from the vertebrate paleontology community.
One might say that "stage of evolution" has undeservedly received a bad following in
the press, especially when its importance in age assignment of terrestrial mammal
faunas is acknowledged.
In agreement with the critics, however, it must be emphasized that age deter-
mination based ouly on "stage of evolution" is hazardous at best. On the other hand,
when mammal faunas have been well studied, as is generally true in Europe, with
several lineages (e.g., horses, primates, theridomyids, cricetids, and gomphotheres)
well known and widely distributed, the "evolutionary grade" or "stage of evolution"
within each of these lineages is generally corroborative and age assignment of a par-
ticular faunal assemblage is usually straightforward. In those instances, most verte-
brate paleontologists would place as much confidence in relative age assignment based
on "stage of evolution" as they would on stratigraphic superposition. In fact, the
Mammal Neogene (MN) zones proposed by P. Mein and co-workers for European
mammal faunas about 15 years ago, represent a synthesis of some of these lineages
(plus appearances of immigrant taxa). The European MN zonation has been widely
applied and is currently the most reliable chronologie framework for Neogene mammal
faunas of Europe. A number of problems have been identified in recent years, both in
the assignment of individual faunas to specific MN zones and in the characterization
of some MN zones. These problems are inevitable in any chronologie system; they
represent growth through scientific testing. The contribution by P. Mein (Chapter 6)
is a revision of the European MN zonation, in response to those problems. The MN
zones must be tested, criticized, and revised so that vertebrate paleontologists will
develop confidence in their use.
Perhaps the most reliable and "irreversible" demonstrations for stage of evolu-
tion involve the height of crown in herbivorous mammals such as horses, bovids, and
numerous small mammals (e.g., cricetid and microtine rodents). Certainly, the
observed trends of increase in crown height and steepening of crowns, reduction of
roots, development of cementum and development of dentine tracts in these mammals
3
(illustrated by Fejfar and Heinrich, Chapter 7, figure 5) from incremental, non-
reversible grades, serve as excellent examples of the "stage of evolution" concept.
Mammal Dispersal Events
Dispersal events are the third basic strategy utilized by vertebrate paleontol-
ogists for chronological age assignment. Application of radiometric and paleomag-
netic data to biostratigraphic data permit the development of a high-resolution
chronostratigraphic framework where thick and relatively complete stratigraphic
sections occur. This high-resolution chronologie framework has created new avenues
for refined analysis of both short-term or local stratigraphic events, such as storm
beds, and long-term, regional or global stratigraphic events, such as meteorite
impacts. Identification of these stratigraphic signals have revitalized stratigraphic
geology, wherein specific isochrons or time lines can be traced across a depositional
basin, or across continental areas (Kauffman, 1988). Mammal dispersal events are the
best known and most widely used biostratigraphic events applicable to terrestrial
depositional systems. Mammal dispersal events have been used by vertebrate paleon-
tologists for many years to mark boundaries of biochronologic units. For example, the
Astaracian/Vallesian or MN 8/9 boundary is broadly recognized as the Hipparion
dispersal event, and in North American the Hemingfordian/Barstovian boundary is
broadly recognized as the Gomphotherium dispersal event. Prior to the advent of
"event stratigraphy" these boundaries were considered synchronous, and the appear-
ance of a genus in the region of its origin was considered roughly equivalent to its
appearance in regions where it had dispersed. High-resolution chronology demon-
strates the timing and duration of these dispersal events (e.g., Lindsay et al., 1980,
1984) which provide data for interpreting the dynamics of faunal interchange and
possible trigger mechanisms that initiate dispersal events. We need to learn more
about sea level change in the area where migration occurred, and climatic control of
eustatic sea level, as well as the ancestry of immigrants and timing of dispersal events
in order to analyze the dynamics of faunal interchange. The point is, none of this is
meaningful without a high-resolution chronologie framework.
The most widely known examples of dispersal events come from the history of
horses, eloquently characterized by Simpson (1961) and more recently reviewed by
MacFadden (1985). These horse dispersal events are but a few of the currently known
(or suspected) dispersal events between Eurasia and North America. Timing and
duration for most dispersal events are poorly known but are actively being resolved, as
discussed in this volume. Appearance of the horse Anchitherium in Europe was
initially intended as a guide to the beginning of Aragonian Stage but subsequent work
by Daams and Freudenthal (1977) showed that other fossil "guides" are more reliable,
especially in Spain (see Chapter 4 by Daams and Freudenthal). Similarly, the appear-
ance of Cormohipparion and Hipparion was calibrated and widely used as the European
Hipparion Datum at 1Z.5 Ma (Berggren and Van Couvering, 1974) but later work
demonstrated the appearance of "Hipparion" in southern Asia about 3 Ma later, at
9.5 Ma (see Chapter 34 by Barry and Flynn). The study of Sen (Chapter Z9) has placed
the appearance of "Hipparion" in the Mediterranean area at about 11.5 Ma, narrowing
the difference between these appearances but still suggesting a sequential dispersal of
"Hipparion," or a large gap in the resolution of Cenozoic mammal chronology. The
contributions by Tassy (Chapter 16) and Antunes (Chapter 17) address the
Proboscidean dispersal event; both of these authors conclude that there were several
Proboscidean dispersal events. The contribution from Masini and Torre (Chapter 9)
addresses the diachronous appearance of several mammal immigrants to the Italian
peninsula during the later Cenozoic.
Application of mammal dispersal events is limited by knowledge of mammal

faunas on a global scale. Dispersal events demand a global perspective; otherwise,
immigrants could not be recognized. As the faunas of Asia and Africa become better
known and more accurately calibrated, dispersal events will be constrained and will
test the possible synchrony of dispersal events on separate continents. Barry and
Flynn (Chapter 34) list 133 calibrated faunal changes in the Siwalik sequence of
Pakistan. This is a very positive step toward identifying mammal dispersal events;
4
however, few other continental areas have thick, well exposed, fossiliferous sequences
comparable to those found in Pakistan.
Knowledge of mammal faunas on a global scale will also test the geographic
limit of climatic and tectonic events suspected to have global significance. The
reality of climatic and tectonic events triggering biologic changes all over the world is
presently open to question; mammal dispersal events are one of the best measures for
evaluating the geographic limit and magnitude of climatic events suspected to have
global effects. For instance, widespread cooling and lowering of sea level at about
Z.4 Ma had a profound effect on mammals and could have triggered the dispersal of
mammals between continents. It seems as though the dispersal of mammals between
North and South America, the Great American Interchange, was almost coincident
with the Z.4 Ma lowering of sea level (Marshall, 1985). However, the dispersal of
elephants from Africa to Europe and Asia was probably slightly earlier than the Z.4 Ma
cooling and lowering of sea level. Similarly, the "global" effect of the emerging
Himalaya Mountains may have triggered one or more late Miocene mammal dispersal
events. The tools are at hand to evaluate the synchrony (or diachrony) of many
dispersal events; however, assembling the appropriate data has rarely been given high
priority. Synthesis of this information can best be accomplished by vertebrate paleon-
tologists as it always involves taxonomic as well as chronologie data. There is
currently too much speculation about "global" events in earth history and not enough
hard data on the timing of immigrant appearances. However, this problem is being
corrected, as seen in numerous contributions to this volume.
BIOCHRONOLOGY, BIOSTRATIGRAPHY, CHRONOSTRATIGRAPHY,

MAMMAL STAGES, MAMMAL AGES, AND CHRONOZONES
The following discussion is intended to clarify some widely used terms and their
use in mammal chronology.
Biochronology can mean the study of any temporal aspects of life, or the study
of life forms with respect to time, or the timing of life forms, or simply the study of
biochrons. A biochron was defined by Williams (1901) as "a time unit whose measure is
the endurance of organic characters." The International Stratigraphic Guide (Hedberg,
1976) characterizes a biochron as the time represented by a biozone, and a biozone is
presented as a general term for any kind of biostratigraphic unit. The latter definition
is rather vague and liable to confuse paleontologists or anyone else who tries to distin-
guish between a biochron, a biozone, or a biostratigraphic range zone. In practice, all
units with the term or suffix "zone" (e.g., range zone, teilzone, assemblage zone, or
biozone) are best applied to fossils in association with a body of rock; that is, for
biostratigraphic associations. Specifically, a biozone is a general expression for any
type of biostratigraphic unit; a range zone should indicate the total stratigraphic
range of some fossil taxon; a teilzone should indicate the local stratigraphic range of
some taxon; and an assemblage zone characterizes the stratigraphic interval of a
particular group of fossils. Note that none of these expressions designate a unit of
time. Technically, when we want to express the time represented by a biostrati-
graphic unit we change it to a chronostratigraphic unit and the ter:m "zone" to "chron,"
as in biochron, range (range chron seems redundant), and teilchron. The time repre-
sented by an assemblage zone is a chronozone, or part thereof.
The term biochronology was rarely used prior to about 1970 when the application
of radiometric dating became widespread in geology, and the distinction was made
between radiochronology and biochronology as different aspects of geochronology (see
Berggren and Van Couvering, 1974). Berggren and Van Couvering (op. cit., p. 6)
suggested application of the term "biocbron" for units of geologic time that are based
on paleontologic data without reference to lithostratigraphy or rock units. This
follows logically from Article ZZ(h) in the 1961 and 1970 North American Codes of
Stratigraphic Nomenclature (published by the AAPG) and the International
Stratigraphic Guide (Hedberg, 1976). However, biochron is not adequately defined in
any of those stratigraphic guides.
5
Biostratigraphy can indicate the sequence of life forms that were preserved in
rock, or the description of rock based on life forms. The International Stratigraphic
Guide (Hedberg, 1976) characterizes biostratigraphy as "the element of stratigraphy
that deals with the remains or evidences of former life in strata, and with the organi-
zation of strata into units based on their fossil content." Biostratigraphy has a long
and well established history as the paleontological aspect of sedimentary geology.
Assemblage zones are the fundamental biostratigraphic unit and superposition is the
main or fundamental strategy for ordering units in biostratigraphy.
Biostratigraphy, in contrast to biochronology, is not a measure of time; it is a

characterization of stratified rocks based on the fossil content within those rocks.
Local stratigraphic sections will always record stratigraphic gaps or hiatuses as well
as faunal change. Therefore, boundaries of assemblage zones very likely represent
gaps or hiatuses. The best way to fill in those gaps is to develop additional assemblage
zones in other stratigraphic sections. When several local stratigraphic sections are
assembled into a composite sequence, filling in most if not all of the gaps, chrono-
stratigraphic units can be identified. It is implicit that any local stratigraphic section
and the assemblage zones developed therein will include temporal gaps and missing
faunal elements. Prior to the development of isotopic dating and magnetostratig-
raphy, stratigraphic gaps were identified or eliminated by constructing concurrent-
range zones, that is by developing a more robust fossil record through replicate strati-
graphic sections.
Many of the contributors to this volume apply the terms biostratigraphy and/or
biozones for sequences of faunas whose ordering is based on stage of evolution rather
than stratigraphic superposition. If mammal faunas are ordered by stage of evolution,
the terms biochronology and/or biochrons are more appropriate to describe the
chronologie ordering and sequence. For instance, Mammal Neogene (MN) zones, as
developed and used in Europe, are ordered primarily on the basis of evolutionary grade
rather than stratigraphic superposition. Therefore, MN zones are best termed bio-
chronologic units.
Chronostratigraphy, on the other hand, is a relationship of time and rock

sequences. The distinctions between biostratigraphy and chronostratigraphy are
significant and should be emphasized. Simply stated, time units cannot overlap and
they must include all temporal segments. Biostratigraphic units can record gaps;
chronostratigraphic units cannot. Biostratigraphic units, often representing ecological
divisions or incomplete sections, can become chronostratigraphic units only when
replicate sections and a more robust fossil record provide evidence that all the facies
are represented and all the gaps have been filled.
Another important characteristic of chronostratigraphic units is that boundaries

of adjacent units must be isochronous. In most instances isochrony of boundaries is
only implied rather than demonstrated; the level of confidence that "all of included
time is represented, without gaps or overlap" is implicitly a condition of boundary
isochrony.
Mammal stage has been applied as a chronostratigraphic unit (e.g., Vallesian

Stage) based on European mammal evolution. This usage implies that the chronologie
interval representing Vallesian Stage has been established from a biostratigraphic
framework, using stratigraphic superposition, that the lower boundary of the unit is
well defined and reasonably isochronous, and that the unit has been tested to the
satisfaction that "all of included time is represented, without gaps or overlap." I
suspect that most European mammal stages have not been adequately tested for
completeness, and that the lower boundaries are not well defined nor tested for
isochrony and overlap. Perhaps the European mammal stages are reliable chrono-
stratigraphic units; the ultimate test of their reliability is how long they will be found
useful.
A type section is another implicit requirement for a mammal stage, because

chronostratigraphic units should be based on stratigraphic superposition. However,
6
designation of a type section, as for the Vallesian Stage, does not "create" a mammal
stage. The procedure for establishing a mammal stage should be, first, to establish
assemblage zones, then to test these assemblage zones by developing more assemblage
zones in other sections. Gradually, a chronologie framework will evolve, as all the
gaps are filled. Designation of a type section, or type sections, should be the last step
in creating a mammal stage, after the most complete and best represented strati-
graphic sequences have been identified. Mammal stages can be developed but they
should be tested prior to designation as chronostratigraphic units, to demonstate their
reliability and limits.
The phrase mammal stage could also be applied to a land mammal age that had
been well established and widely recognized, but which had subsequently been placed
in a biostratigraphic framework, and thereby transformed into a chronostratigraphic
unit. In this example, the interval of time has not changed; however, it has been more
securely placed by demonstration of superposition. This scenario has occurred twice
in North America, wherein Savage (1977) proposed Wasatchian Stage for the
Wasatchian land mammal age, and Rose (1981) redefined Clarkforkian land mammal
age, nominating it as a chronostratigraphic unit. It should also be pointed out that the
distinction between these units, as land mammal ages or stages, is insignificant and no
increase in resolution or accuracy in correlation has been detected subsequent to these
changes. Certainly, demonstration of superposition for biochronologic units is benefi-
cial; however, it is not essential.
Mammal ages are most commonly used in North America where they were devel-
oped over the last 50 years. The development of North American land mammal ages is
reviewed by Lindsay and Tedford (Chapter 37) and will not be repeated here.
Berggren and Van Couvering (1974, pp. 91-118) noted that North American land
mammal ages and European mammal stages were in reality biochrons. Woodburne
(1987, p. 1) also emphasized that North American land mammal ages are not equiva-
lent to biostratigraphic units, but come closest to the concept of biochrons as
proposed by Williams in 1901. Woodburne inferred that vertebrate paleontologists in
North America should identify their terrestrial chronologie system as a biochronology
to distinguish it from the marine chronologie system of biostratigraphy. Woodburne
did not mean to diminish the significance of biostratigraphic applications; he is a
strong advocate of biostratigraphic methods. His point is that the intrinsic nature of
terrestrial deposits is better suited for the development of biochrons rather than
assemblage zones, and the resulting chronologie framework is closer to biochronol-
ogy. Another important point made by Woodburne is that the level of reliability and
resolution in the current North American land mammal ages is comparable to the
reliability and resolution in the best marine chronostratigraphic systems.
A final comparison should be made between the mammalian chronologie frame-

work of Europe and North America, both of which are based primarily on biochrono-
logic concepts. The European chronology has a more robust, better studied taxonomic
base but a weaker, less complete stratigraphic base. In contrast, the North American
chronology has a broader stratigraphic base but a weaker, less robust taxonomic
base. Both chronologie frameworks require repeated testing for refinement and
confidence, by both the stage of evolution and superposition strategies. Ideally, both
strategies will be applied whenever feasible; in reality, the stage of evolution strategy
will be applied more readily in testing the European chronology, and superposition
strategy will be applied more readily in testing the North American chronology.
Chronozones were invented with the 1970 North American Code of Stratigraphic
Nomenclature. The term has subsequently been added to the International
Stratigraphic Guide (Hedberg, 1976) and the 1983 North American Stratigraphic Code
(Bulletin, American Association Petroleum Geologists, v. 67, 1983). Chronozones were
conceived as low-ranking chronostratigraphic units equivalent in time span to a bio-
stratigraphic unit or other zone. They have never been applied widely, and their
application is likely to be loose or confusing. Implicitly, chronozones should be
(1) stratal units, with a designated stratotype, and (2) boundaries of chronozones are
7
isochronous, by definition. Chronozones might be applicable as subdivisions of
mammal stages. However, it has never been clear whether chronozones are applicable
as biochronologic units. If so, the Mammal Neogene (MN) biochrons approach the
concept of chronozones. MN zones (or biochrons) are frequently used by European
vertebrate paleontologists as though they were chronozones. However, the lower
boundary of an MN biochron would have to be defined and demonstrated isochronous,
and its reliability demonstrated through use for it to qualify as a chronozone. I
believe these criteria probably can be met, and that MN biochrons could be designated
MN chronozones. However, they cannot be assumed chronozones by definition; they
can become chronozones only through demonstrated use, or testing.
To summarize, chronologie systems that require the strategy of superposition for

ordering the chronology are best termed biostratigraphic systems. Chronologie
systems that utilize stage of evolution (usually because they lack a stratigraphic
foundation) as the primary strategy for ordering the chronology are best termed
biochronologic systems. It follows that the chronologie framework developed by
vertebrate paleontologists for mammal evolution in both North America and Europe
are biochronologic systems. However, it should be emphasized that biostratigraphy
and the strategy of superposition are essential for application of isotopic dating and
magnetostratigraphy in the calibration and correlation of mammalian chronology.
TOWARD A GLOBAL CHRONOLOGY FOR MAMMAL EVOLUTION
Great strides were made during the mid-1970s toward development of a global
chronology for mammal evolution. In part, this was triggered by publication of The
Late Neogene by W.A. Berggren and J.A. Van Couvering in 1974. Berggren compiled
the marine biostratigraphic record, Van Couvering compiled the mammalian bio-
chronologic record, and together they synthesized a late Neogene global chronologie
framework, emphasizing interregional correlations and summarizing climatic events
using glacial and floral data to supplement the biostratigraphic data. Berggren and
Van Couvering's synthesis came on the heels of new isotopic age determinations from
terrestrial deposits and magnetic polarity data from deep sea cores; it provided
reliable and hard data that would allow paleontologists and geologists in widely
separated areas to test the synchrony or sequence of biologic and geologic events from
a global perspective.
European Mammal Sequences
During 1975 the foundations for the current framework of European mammal
chronology were laid. In April of 1975 the first international symposium on European
mammal chronology was held at Munchen (Fahlbusch, 1976). During September of
1975, P. Mein presented an expanded compilation of Mammal Neogene (MN) zones at
the meeting of the Regional Committee on Mediterranean Neogene Stratigraphy in
Bratislava. The resulting chronologie framework was based on a sequence of refer-
ence localities, many of those reference localities were identical with reference local-
ities given for the MN zones by Mein, and the MN zones were incorporated into the
resulting European Neogene mammal chronology.
Chapters Z through 9 address the broader chronologie framework and more

general problems of European mammal chronology. The contribution by Steininger et
al. (Chapter Z) is a new synthesis of European chronologie systems, combining data
from both marine and terrestrial deposits. Note that both biostratigraphic and bio-
chronologic sequences are given for European Neogene mammal faunas. This chapter
is followed by a review of the synthesis (and publication) of Paleogene biozonation for
European mammal faunas by Schmidt-Kittler (Chapter 3), who edited that publica-
tion. Chapters 4 and 5 summarize biostratigraphic syntheses that apply to European
mammal chronology. Daams and Freudenthal (Chapter 4) summarize the extensive
and detailed biostratigraphic work they have undertaken in Spain since 1976. Every-
one agrees that Spain has the best stratigraphic sequence for developent of a mammal
8
chronology in Europe. The next chapter by Sumengen et al. (Chapter 5) presents new
biostratigraphic results from Turkey. Turkey, bordering Europe, has a good Tertiary
sequence and is ideally suited to identify biogeographic limits of Europe and Asia.
Chapters 6 and 7 present the framework for applying the stage of evolution concept in
European Neogene chronology. Chapter 6 is a revision and update of Mammal
Neogene (MN) zones by P. Mein. Fejfar and Heinrich (Chapter 7) define biochrono-
logic units (zones and superzones) for Europe, based on muroid rodents, widely recog-
nized as the most abundant, diverse, and well studied groups of mammals during the
Neogene of Europe. Differences between MN zones of Mein and muroid zones (and
superzones) of Fejfar and Heinrich are not obvious. With further testing and possible
revision, these units will likely be combined into broader based and more precisely
defined biochronologic units. Chapter 8, by Guerin, addresses problems of developing
a comprehensive biochronologic framework for all of Europe, focusing on Quaternary
mammal faunas. In extending the Mammal Neogene zones, where does one draw the
line, especially when personal bias, taxonomic documentation, and nationalistic
preference enter the equation? Chapter 9, by Masini and Torre, points to large
mammal dispersal events as a key for recognition of boundaries and subdivision for
Villafranchian land mammal age. Note that the Villafranchian of Masini and Torre,
the Villanyian of Fejfar and Heinrich, and MNQ 16 and 17 of Guerin are all the same,
or are they?
Together, this and the following eight chapters provide an overview of European
mammal chronology, and background for the following chapters that address more
specific problems of European Neogene mammal chronology.
The contributions of Hugueney and Ringeade, Ginsburg, Engesser, Heissig,

Kowalski, and Fejfar summarize the mammal faunal sequence from important but
restricted areas of Europe. Hugueney and Ringeade (Chapter 10) provide a compre-
hensive review of the rodent sequence in the Aquitaine Basin where the concept of
Agenian land mammal age (early Miocene) was defined. Ginsburg (Chapter 11) reviews
recent collecting in the Loire Basin of France and presents revised faunal lists for
many critical faunas of that area. Both Hugueney and Ringeade's and Ginsburg's
studies can be viewed as current "stage of the art" for European biochronology, com-
bining biostratigraphy, stage of evolution, and dispersal events to order the local
sequence of mammal faunas. Engesser (Chapter 12) reports on new and scant remains
of land mammals from Switzerland, in deposits that were formerly thought entirely
marine, lacking terrestrial mammals. Engesser places 12 mammal sites into four MN
zones, establishing superpositional and marine stratigraphic relationships for these
mammal faunas. This work is ongoing and promises to strengthen correlation between
MN zones and marine biostratigraphic chronologies. Heissig also reports (Chapter 13)
new rodent finds from southern Germany which fill an unidentified gap in the middle
Miocene mammal sequence of that area. Kowalski (Chapter 14) and Fejfar (Chapter
15) summarize the Cenozoic mammal faunas known from Poland and Czechoslovakia,
respectively, ordering these faunas in the chronologie framework recognized over
other parts of Europe. These Polish and Czechoslovakian faunas are key references
for correlation of eastern European and Asian faunas.
Contributions by Tassy, Antunes, Bemor et al., Koufos, and Azzaroli address

large mammal dispersal events and their resolution for European Neogene
Chronology. Tassy (Chapter 16) and Antunes (Chapter 17) review and summarize the
Proboscidean Datum Event, noting there are several proboscidean dispersal events but,
more important, the probability of several dispersal routes for these mammals.
Bemor et al. (Chapter 18) summarize the record of hipparionine horses in Europe,
along with the presumed ancestors of those horses in North America. Note that Sen,
in Chapter 29, also addresses the European Hipparion datum, but Sen emphasizes the
timing of Hipparion dispersal whereas Bernor et al. emphasize the systematics of
Hipparion horses. Koufos (Chapter 19) examines the record of hipparionine horses
from northern Greece, and places these records in a biochronologic framework.
Azzaroli (Chapter 20) reviews the European record of Equus, emphasizing the distribu-
tion of Equus species in late Neogene and Quaternary faunas of Europe.
9
Contributions by Moya-Sola and Agusti, by Agusti, by Storch and Fejfar, and by
Pickford address biogeographic problems of European Neogene mammal chronology.
Moya-Sola and Agusti (Chapter Z1) summarize mammal associations recorded from
Miocene deposits of Spain and interpret these associations relative to major events in
earth history. Agusti (Chapter ZZ) examines the biogeographic affinity of rodents
recorded from eastern Spain to identify biogeographic provinces and their limits
during the Miocene. He finds the Miocene associations have changed significantly,
inferring a shift of zoogeographic boundaries. Storch and Fejfar (Chapter Z3) describe
a small mammal fauna from northern Germany, finding that its affinity with faunas of
Asia is as strong as the affinity with more temperate European faunas. Pickford
(Chapter Z4) examines the factors that limit biogeographic distribution of land
mammals, noting changes in distribution patterns that occurred during the Miocene; he
concludes that the boundary between the Palearctic and Ethiopian biogeographic
provinces has probably shifted significantly during and after the Miocene. All of these
studies (Chapters Z1 through Z4) point to the need for caution when identifying
Neogene boundaries of biogeographic provinces.
Contributions by Alpagut and Andrews, by Alvarez-Sierra et al., by Bonifay, and

by Andrews address ecological factors that influenced the distribution and preserva-
tion of mammals during the Neogene. Alpagut and Andrews (Chapter Z5) identify the
community structure of the Pasalar faunas in Turkey and compare it with the com-
munity structure of modern faunas. They conclude the Pasalar community most
closely resembles the subtropical, semi-deciduous forests of India, with summer
monsoonal rainfall. Alvarez-Sierra et al. (Chapter Z6) examine the mammal record of
Spain for evidence of non-biologic factors that leave a distinctive signature or over-
print on the record. They record abrasion and corrosion in samples of eomyid and
glirid rodents, noting significant variance. They conclude that differences in abrasion
of eomyids and glirids is evidence that these small mammals were not concentrated by
the same mechanism (i.e., owl predation). Bonifay (Chapter Z7) examines late
Cenozoic mammal sequences of western Europe from both a taxonomic and geographic
perspective. She notes that major intervals of mammal turnover (e.g., Villafranchian
and Quaternary) do not coincide with major climatic events in the marine record.
Andrews (Chapter Z8) reviews taphonomic factors that concentrate bone in small
mammal faunas. Small mammals, especially rodents, are very important members of
European Neogene mammal faunas; knowledge of factors that control their preserva-
tion and high concentration are critical for both learning where to find these concen-
trations and how to interpret their record for both ecology and chronology. All of
these studies (Chapter Z5 through Z8) emphasize the need for understanding paleoeco-
logical biases when interpreting chronological sequences of mammal faunas.
Contributions by Sen, Opdyke et al., and Langereis et al. demonstrate the

application of magnetostratigraphy to problems in European Neogene mammal
chronology. Magnetostratigraphy is a very powerful tool for sequential ordering of
mammal faunas. Sen (Chapter Z9) addresses the "Hipparion Datum" in the circum-
Mediterranean area. This is a key biochronologic marker for middle Miocene faunas of
Europe. Sen notes the appearance of "Hipparion" is slightly diachronous in the
Mediterranean area, occurring in chron 10 and possible in chron 11; he concludes that
the Mediterranean "Hipparion Datum" is considerably older than the Siwalik
"Hipparion Datum" which is in chron 9. Opdyke et al. (Chapter 30) present magneto-
stratigraphic sections from late Miocene terrestrial sections in the Cabriel Basin of
Spain. Their correlation shows the placement of MN 1 Z and 13 faunas within the
interval of magnetic chrons 7-5. Langereis et al. (Chapter 31) present magnetostrati-
graphic sections from stratigraphic sequences in Turkey, sequences collected and
reported in the contribution by Sumengen et al. (Chapter 5). These two contributions
(Chapters 5 and 31) are pioneering efforts to establish a high-resolution chronology in
this key geographic area between Europe, Asia, and Africa. These three magnetic
polarity studies show promise for further application of magnetostratigraphy in Europe
and surrounding areas. Additional studies will be needed to limit the entire Neogene
European mammal sequence relative to the magnetic polarity time scale. The paleo-
magnetic studies reported here provide initial hypotheses for the limits of MN zones
9-13 relative to magnetic chrons 5-10; other stratigraphic sections must be found and
10
sampled during the coming years, to test these hypotheses and confirm or revise the
chronologie framework. Calibration of European mammal ages or stages is not too far
in the future.
Asian Mammal Sequences
Contributions from Qiu, Barry and Flynn, plus Jacobs et al. address Neogene
mammal faunas of Asia and their relationship to European Neogene mammal faunas.
Qiu (Chapter 3Z) presents an expanded sequence of Neogene mammal faunas in China,
ordering these faunas in an informal biochronologic framework, placed (broadly)
relative to the European mammal chronology. During the Cenozoic, central Asia was
one of the largest emergent continental areas and probably a center for evolution of
many mammal groups. Central Asia, especially China, has one of the best records of
Neogene mammals, based on the abundance of terrestrial deposits there. Great
strides have been and are being made to document the record of Neogene mammals in
China. Qiu summarizes the history of fossil mammal collecting in China, and
organizes the Chinese mammal record in a sequentially numbered framework. This
framework will prove very useful for correlations with Europe, and will provide a
foundation to build upon. Barry and Flynn (Chapter 33) and Jacobs et al. (Chapter 34)
address the Siwalik record of Pakistan in southern Asia. The Siwalik record is one of
the most complete and best calibrated terrestrial sequences presently known. Barry
and Flynn discuss biostratigraphic events and their calibration in the Siwalik
sequence. Jacobs et al. focus on the abundant Siwalik record of muroid rodents,
reflecting on how the Siwalik record differs from the European record of similar
rodents; they conclude that the modern diversity of muroid rodents results from
multiple dispersal events from central Asia since the middle Miocene.
African Mammal Sequences
Vertebrate history in Africa was comprehensively reviewed by Maglio and Cooke

(1978), in which a summary of the sequence of African mammal faunas was presented
(on p. Z9). At about the same time, Szalay and Delson (1979) reviewed the fossil
Primate record, including a set of undefined African land mammal ages (Fayumian,
Rusingan, Ternanian, Ngororan, Lothagamian, and Rodolfian). These same African
land mammal ages were applied and amended (replacing Rodolfian with Langebaanian
and Makapanian) in the encyclopedic review of Cenozoic mammals by Savage and
Russell (1983). Pickford (1986) grouped and ordered a number of mammal sites
(mostly Miocene, late Pliocene, and Pleistocene) into faunal "sets" to demonstrate the
sequence of mammal faunas in western Kenya. Thus, for part of the Cenozoic (espe-
cially the Miocene) a sequence of African vertebrate faunas has been named and
designated as "land mammal ages" or similar units, although these units and their
boundaries have not been formally defined.
R. Savage (Chapter 35) reviews the early Neogene record of Africa and describes
the faunal composition of the Gebel Zelten fauna. The Gebel Zelten and similar
Orleanian-equivalent faunas from Africa are critical for interpreting European-
African mammal dispersal events near the Proboscidean datum. Savage places these
faunas relative to the European MN zones.
North American Mammal Sequences
Lindsay and Tedford (Chapter 36) review the development of biochronology in

North America and Europe. A comprehensive summary of North American
mammalian biochronology (Cenozoic Mammal Faunas of North America), edited by M.
Woodburne, was published in 1987. It has taken three generations of vertebrate
paleontologists (about 40 years) to attain the accuracy, resolution, and confidence
presently recognized for North American land mammal ages. In contrast, develop-
ment of a chronologie framework for European mammal faunas, free from a founda-
tion based on marine stratigraphy, has existed little more than 15 years, spanning only
one generation of vertebrate paleontologists. Biochronology of European mammal
faunas is still "maturing" relative to North American mammal biochronology, with
great steps taken in European mammal chronology during recent years.
11
SOGth American Sequences
Neither South American nor Australian mammal faunas are discussed further in
this volume. These notes on mammal chronology in South America and Australia are
presented to complete the picture of global mammal history.
Land mammal ages were suggested for South America by Patterson and Pascual
(1968) and by Simpson (1971, p. 107). These chronologie units were derived primarily
from the work of Simpson (1940), in which he proposed a series of terrestrial "stages"
for South America. Addition (e.g., ltaboraian), revision (e.g., Deseadan and
Colhuehuapian), and calibration of these South American land mammal ages were
presented by Marshall (1985) and MacFadden et al. (1985). Thus, during the last ten
years great strides have been made toward chronologie resolution of terrestrial
deposits in South America.
Australia
A sequence of mammal faunas from Australia was published by Stirton, Tedford,

and Woodburne (1968). In that study faunal units were ordered by stage of evolution,
or (when possible) superposition. The resulting sequence was placed in a framework of
Australian marine stages. Subsequent work (summarized by Woodburne et al., 1984)
has increased that data and provided limited calibration, based on intercalation with
volcanic units. Woodburne et al. (1984, figure Z) placed the expanded Australian and
New Guinea mammal chronology in the same framework of marine stages. Boundaries
of the Australian marine stages, as well as the mammal faunas, are still poorly
defined.
Summary
Fahlbusch and Mein (Chapter 37) summarize the prec1s1on and resolution of
European Neogene mammal chronology, and discuss problems that must be addressed
to improve the European chronologie framework.
A comprehensive, well-calibrated record of mammals on all continents is a long-

term goal of vertebrate paleontologists. As we approach that goal, new insight on the
dynamics of fossil populations, shifting biogeographic province boundaries, and the
magnitude of climatic events in earth history are coming to light. Knowledge of
mammal history is now in a transitional state; developed more completely in North
America and Europe, with much recent progress in Asia, South America, and
Australia. Each continent and region has a different set of problems that require
special solutions. Many mammal groups were good travelers whose partial history on
separate continents links the provincial chronologies into a dynamic and unified
history. When traced back in time, the faunal provinces of today might fragment or
merge to show periods of lesser or greater affinity with adjacent provinces. A major
goal is to discern these separate histories in a chronological system that will accurate-
ly depict the major events in each faunal province. Another goal is to show how major
events within one province relate to major events in the others.
During the last fifteen years vertebrate paleontologists have made great
progress toward resolving stratigraphic-taxonomic problems and filling chronologie
gaps in mammal evolution on all the major continents, but especially in Europe. The
picture of mammal evolution is much clearer now than it ever has been, but we are a
long way from achieving a global perspective of mammalian evolution.
ACKNOWLEDGMENTS
I am deeply indebted to my colleagues V. Fahlbusch and P. Mein for their help in

organizing the Schloss Reisensburg NATO Workshop; and to all the participants in the
Workshop my sincere thanks for stimulating discussions, congenial camaraderie, plus
their thoughtful and diligent suggestions toward developing a better chronologie
framework for European Neogene mammal history.
12
This contribution was conceived after the Workshop, in response to some of the
discussions. John Barry, Larry Flynn, Lou Jacobs, and Dick Tedford read an earlier
draft and suggested numerous improvements. To each of them, my sincere thanks.
REFERENCES
Berggren, W.A. and Van Couvering, J.A., 1974. The late Neogene, biostratigraphy,
geochronology, and paleoclimatology of the last 15 million years in marine and
continental sequences. Elsevier Publishing Co., Amsterdam, Z16 p.
Daams, R. and Freudenthal, M., 1981. Aragonian: The stage concept versus Neogene
mammal zones. Scripta Geologica, v. 6Z, p. 1-17.
Daams, R., Freudenthal, M., and Weerd, A. Van de, 1977. Aragonian, a new stage for
continental deposits of Miocene age. Newsletter Stratigraphy, v. 6, p. 4Z-55.
Fahlbusch, v., 1976. Report on the International Symposium on Mammalian Stratig-
raphy of the European Tertiary. Newsletter Stratigraphy, v. 5, p. 16Q-167.
Hedberg, H.D. (ed.), 1976. International Stratigraphic Guide. John Wiley & Sons
Publishers, New York, ZOO p.
Kauffman, E.G., 1988. Concepts and methods of high-resolution event stratigraphy:
Annual Review Earth and Planetary Science, v. 16, p. 605-654.
Lindsay, E.H., Opdyke, N.D., and Johnson, N.M., 1980. Pliocene dispersal of the horse
Equus and late Cenozoic mammalian dispersal events: Nature, v. Z87, p.
135-138.
Lindsay, E.H., Opdyke, N.D., and Johnson, N.M., 1984. Blancan-Hemphillian land
mammal ages and late Cenozoic mammal dispersal events: Annual Review Earth
and Planetary Science, v. 1Z, p. 445-488.
MacFadden, B.J., 1985. Patterns of phylogeny and rates of evolution in fossil horses:
Hipparions from the Miocene and Pliocene of North America: Paleobiology, v.
11, p. Z45-Z57.
MacFadden, B.J., Campbell, K.E., Jr., Cifelli, R.L., Siles, 0., Johnson, N.M., Naeser,
C.W., and Zeitler, P.K., 1985. Magnetic polarity stratigraphy and mammalian
fauna of the Deseadan (late Oligocene-early Miocene) Salla beds of northern
Bolivia: Jo~al of Geology, v. 93, p. ZZ3-Z50.
Maglio, V.J. and Cooke, H.B.S. (eds.), 1978. Evolution of African Mammals. Harvard
University Press, Cambridge, Mass., 641 p.
Marshall, L.G., 1985. Geochronology and land mammal biochronology of the Trans-
american faunal interchange, in Stehli, F.G. and Webb, S.D. (eds.), "The Great
American Biotic Interchange," p. 49-85.
Mein, P., 1975. Resultats du Groupe de Travail des Vertebres, in Report on Activity
of the RCMNS Working Groups (1971-1975), Bratislava, p. 78-81.
Patterson, B. and Pascual, R., 1968. The fossil mammal fauna of South America.
Quarterly Review of Biology, v. 43, p. 409-451.
Pickford, M., 1986. Cainozoic paleontological sites of western Kenya: Munchner
Geowissenschaftliche Abhandlungen, Reihe A., v. 8, p. 1-151.
Rose, K.D., 1981. The Clarkforkian land-mammal age and mammalian faunal compo-
sition across the Paleocene-Eocene boundary. Univ. Michigan Papers in Paleon-
tology, no. Z6, p. 1-197.
Savage, D.E., 1977. Aspects of vertebrate paleontological stratigraphy and geochro-
nology, in Kauffman, E.G. and Hazel, J.E. (eds.), "Concepts and Methods in
Biostratigraphy," p. 4Z7-44Z.
Savage, D.E. and Russell, D.E., 1983. Mammalian Paleofaunas of the World. Addison-
Wesley Publ., New York, 43Z p.
Simpson, G.G., 1940. Review of the mammal-bearing Tertiary of South America.
Proceedings, American Philosophical Society, v. 83, p. 649-709.
Simpson, G.G., 1961. Horses. The Natural History Library. Doubleday-Anchor Books,
Garden City, New York, 3Z3 p.
Simpson, G.G., 1971. The evolution of marsupials in South America. Ann. Acad.
brasil. Cienc., (1971), v. 43, p. 103-118.
Stirton, R.A., Tedford, R.H., and Woodburne, M.O., 1968. Australian Tertiary deposits
containing terrestrial mammals. Univ. California Publ. Geological Sciences, v.
77, p. 1-30.
13
Szalay, F .S. and Delson, E., 1979. Evolutionary History of the Primates. Academic
Press, New York, 580 P•
Williams, H.S., 1901. Discrimination of time value in geology. Journal of Geology, v.
9, P• 57Q-585.
Woodburne, M.O. (ed.), 1987. Cenozoic Mammals of North America. Univ. California
Press, Berkeley, 336 p.
Woodburne, M.O., Tedford, R.H, Archer, M., Turnbull, W.D., Plane, M.D., and
Lundelius, E.L., 1984. Biochronology of the continental mammal record of
Australia and New Guinea. Special Publication, South Australia Dept. Mines and
Energy, no. 5, p. 347-363.
14
EUROPEAN NEOGENE MARINE/CONTINENTAL
CHRONOLOGIC CORRELATIONS
Fritz F. Steininger
lnstitut fUr Palaontologie

Universitat Wien
Vienna, Austria
Raymond L. Bernor
Laboratory of Paleobiology
Department of Anatomy
College of Medicine
Howard University
Washington, D.C., U.S.A.
Volker Fahlbusch
lnstitut fiir Palaontologie und historische Geologie

Universitat Miinchen
Miinchen, Germany
WE DEDICATE THIS PAPER TO THE LATE CLAUDIO de GIUU
ZUSAMMENFASSUNG
Eine Korrelationstabelle fUr das Neogen zwischen Magnetostratigraphie und

marinen Planktonzonierungen, den in Verwendung stehenden Mediterranen- und
Zentralen Paratethys-Stufen, den Pollenzonierungen, aber vor allem den Saugetier-
Faunenzonen und Fauneneinheiten sowie neu vorgeschlagener kontinentaler chrono-
stratigraphischer Stufen wird vorgelegt und im Text diskutiert. Eine Datenbank fur
die Korrelation der SM.ugetier-Zonierung ist angefiigt. Die Grenzen der neogenen
Saugetier-Faunen-Einheiten konnen nach dem derzeitigen Kenntnisstand wie folgt
korreliert werden: Agenium/Orleanium (MNZb/MN3-Faunen-Zone) mit der Aquitan/
Burdigal-Grenze (NNl/NNZ); Orleanium/Astaracium-Grenze (MN5/MN6) im Bereich
der Burdigal/Langhe-Grenze (NN4/NN5); Astaracium/Vallesium-Grenze (MN8/MN9) im
Bereich der Serravall/Torton-Grenze (in der NN9); Vallesium/Turolium-Grenze
(MNlO/MNll) im mittleren Torton (in der NNlO); Turolium/Ruscinium-Grenze
(MN13/MN14) mit der Messin/Zanclium-Grenze (in der NNlZ); Oberkante des
Rusciniums (MN15/MN16) im Bereich des tieferen Piacenzium und die Oberkante der
Saugetier-Faunenzone MN16 im Bereich des hb"heren Piacenzium, die Sflugetier-
Faunenzone MN 17 reicht in das tiefere Pleistozb.
European Neogene Mammal Chronology 15

Edited by E.H. Lindsay eta/.
ABSTRACT
An updated correlation is given of the revised Neogene European mammal

chronology with the Neogene marine chronology and the magnetic polarity time scale.
The boundary of the Agenian/Orleanian mammal-faunal units (boundary of

MN2b/MN3 zones) correlates approximately with the Aquitanian/Burdigalian Stage
(NN1/NN2) boundary. The Orleanian/ Astaracian (MN5/MN6) boundary correlates
approximately with the Burdigalian/Langhian Stage (NN4/NN5) boundary; the
Astaracian/Vallesian (MN8/MN9) boundary correlates approximately with the
Serravallian/Tortonian Stage boundary (within NN9); the Vallesian/Turolian
(MN10/MN11) boundary occurs within the middle part of the Tortonian Stage (within
NNlO, resp. between CN8a and CN8b); the Turolian/Ruscinian (MN13/MN14) boundary
coincides approximately with the Messinian/Zanclean Stage boundary and the upper
boundary of the Ruscinian mammal faunal zones (MN15/MN16) correlate approximate-
ly within the Upper Pliocene (resp. the lower Piacenzian Stage). The upper limit of
the MN17 mammal faunal zone occurs within the lowermost Pleistocene.
INTRODUCTION
The NATO Advanced Research Workshop on "European Neogene Mammal

Chronology" organized at Schloss Reisensburg in Bavaria (Germany) by V. Fahlbusch,
E. Lindsay, and P. Mein brought together 48 specialists from 15 countries. This
offered not only the unique possibility to discuss and revise the European mammal
zonation, but also its calibration relative to the marine (mainly planktonic) biochronol-
ogies, the magnetic polarity time scale, and the European Neogene chronostrati-
graphic stage systems.
An ad hoc "correlation group" formed at the Reisensburg meeting concentrated

on the task of correlating the mammal zonation with the marine scale. The correla-
tion group included F. Steininger, L. Flynn, M. Freudenthal, C. de Giuli, 0. Fejfar, plus
the meeting organizers. A first circular with a general concensus proposal and the
structure of a correlation table was sent to all participants of the "correlation group"
by L. Flynn in May 1988. A first draft of a continental/marine-chronology correlation
chart and general text was circulated by F. Steininger in August 1988 not only to the
"correlation-group" members but also to colleagues known for their interest in
Neogene chronology.
Our correlation chart for marine and continental Neogene chronology of Europe
is presented as figure 1. The responses and dates received have been integrated and
are discussed in the relevant sections of the text. The text itself is arranged accord-
ing to the different columns of the correlation chart, with ·additional relevant
comments. The final section is concerned with the data base of the mammal-marine-
continental correlations.
CORRELATION OF MAGNETIC POLARITY TIME SCALE

AND MARINE BIOCHRONOLOGIES
Several years ago, Berggren et al. (1985a,b) published an extensive paper on the
correlation between marine planktonic scales (planktonic foraminifera-calcareous
nannoplankton-radiolarian) respectively, these scales, and the magnetic polarity time
scale. In these two papers magnetic Anomaly 5 is correlated with Chron 11 (respec-
tively Chron C5N), which results in placing the middle/late Miocene boundary, gener-
ally located within Blow's Plankton Zone N15, at 10.5 Ma. This correlation is in con-
trast to earlier papers of Berggren (1984) at 11.8 Ma; Barron et al. (1985a) at 11.8 Ma,
and Ryan et al. (1974) where this boundary is placed at 12 Ma.
Radiometric dates on marine and terrestrial biostratigraphic events as well as

recent magnetic dates at the middle/late Miocene boundary suggest that the correla-
16
0
""'<z
....
0
z
z z
<
::; <
0 ::;
0: 0
0:
~
"'~
......"' z
~
::> z
0
.... z
a: z
~ :!!
VI :!!
VI
~
~
"'::l
<
>
z
<
z
0
C)
<(
"'
<(
MN2b
z
<
z 0
""'z<
z<(
;: "'~
....<(
:>: MN1 0
....
0
z
Fig. 1. Correlation chart for marine/continental Neogene chronology of Europe.
tion between magnetic Anomaly 5 and Chron 11 might have to be revised (Berggren,
pers. comm., August 15, 1988; Sen, pers. comm., October 24, 1988; Baldauf et al.,
1987; Sen, 1986). We therefore follow the earlier and widely accepted correlation of
Ryan et al. (1974), Theyer and Hammond (1974), and Theyer et al. (1978) of magnetic
Anomaly 5 with Chron 9 (resp. Chron C5N).
17
Chron nomenclature follows La Breque et al. (1983). However, recently a so-
called "Systematic Chron Numbering" was proposed by Hailwood (1989).
The correlation between the magnetic polarity time scale and the geochrono-
metric scale follows the recommendations of Berggren (1987) and Berggren et al.
(1985a,b).
The correlation between marine planktonic biozonations, with respect to these

biozonation& and the magnetic polarity scale follows Berggren (1984) and only in parts
Berggren et al. (1985a,b); revisions from these correlations are based on Barron et al.
(1985a), Haq and Takayama (1984), Iaccarino (1985), Martini and MUller (1986), and Rio
et al. (1988).
In terms of the planktonic foraminifera zonations we use here Blow's plankton

zone N4 according to Berggren (1984), Berggren and Miller (1988), Rogl (1981) follow-
ing Iaccarino (1985).
Comments by John Barron (letter dated September 7, 1988) suggest a younger

age for the base of calcareous nannoplankton zone NNll (Chron CN9). This zone
should correlate near the base of magnetic Anomaly 4, at an age of approximately
7.4 Ma (Baldauf et al., 1987; Barron et al., 1985b; Barton and Bloumendal, 1986, in
combination with Lohman, 1986 for DSDP Leg 90).
The assignment of the Messinian/Zanclean stage boundary to a position slightly

below the base of the Thvera magnetic normal subchron in the Gilbert chron (at 4.83
or 4.84 Ma) by Zijderveld et al. (1986) was placed at 4.9 Ma by Channell et al. (1988)
and taken into consideration by Rio et al. (1988) in their redefinitiion and recalcula-
tion of the Pliocene Mediterranean planktonic chronologie framework, which we
follow here.
CHRONOSTRA11GRAPHY: PALEOGENE AND NEOGENE,

MIOCENE AND PLIOCENE, AND THE MEDITERRANEAN
AND CENTRAL PARATETHYS STAGE SYSTEMS
The Paleogene/Neogene (and the Oligocene/Miocene) boundary is defined here in

biostratigraphic terms by the FAD of Globoquadrina dehiscens dehiscens. This bio-
event correlates with Chron C6Br (equals Chron ZZr) with an age of about Z3.Z Ma.
The position of this boundary follows the recommendations and recent results of the
"ICS-Working Group on the Paleogene/Neogene Boundary" (Cati et al., 1981;
Steininger, 198Z; Gelati and Steininger, 1983 as well as Berggren, 1984). The calcare-
ous nannoplankton biozonation cannot be applied as a primary datum because the
boundary between zone NPZ5 and NN1 is inconsistent (see Martini and Muller, 1986).
Specifically, the FAD of Globorotalia kugleri s.str. is complicated by the taxonomic
concept of Globorotalia kugleri s.str. versus the Globorotalia kugleri group, plus
geographic limits of this taxon and relative ease of dissolution of these fossils. These
problems limit the worldwide application of this taxon as a reliable biostratigraphic
marker in the opinion of the P/N-Working Group. In contrast, Globoquadrina dehiscens
dehiscens is more easily recognized, with worldwide range from tropical to temperate
regions, is dissolution resistant, and its first appearance has been shown relatively
synchronous in the Cenozoic oceans. (For further discussion see Berggren et al.,
1985b; Berggren and Miller, 1988; Keller, 1983; Rogl, 1981, and Srinivasan and
Kennett, 1983). Since Globoquadrina dehiscens is already present in the Aquitane-
basin Aquitanian stratotype (Poignant and Pujol, 1979), we follow Iaccarino (1985) in
her correlation of the Aquitanian stage.
The planktonic foraminifera fauna suggests the correlation of the Burdigalian

stratotype in the Aquitane Basin with the base of Blow plankton zone N5. This coin-
cides with the base of the Eggenburgian stage in the Central Paratethys. Also in the
Central Paratethys, the base of the Ottnangian stage correlates roughly with the base
of calcareous nannoplankton zone NN3. The base of the succeeding Karpatian stage is
18
marked by a transgressive event throughout the Paratethys region and in the local
uvigerinid faunas. In the upper part of the Karpatian stage, the FAD of
Globigerinoides bisphaericus is an important planktonic datum as are the NN4 floras
(Rogl and Steininger, 1983).
The early/middle Miocene boundary is defined here by the base of the

Praeorbulina glomerosa s.l.-zone as defined by Iaccarino (1985), with the FAD of
Praeorbulina glomerosa. This event: coincides with the NN4/NN 5 boundary and the
base of the Langhian and Badenian stages, respectively (Bandet et al., 1984; Barron et
al., 1985a; Berggren, 1984 but not 1985a,b; Iaccarino, 1985; Papp et al., 1978). The
early/middle Miocene boundary as drawn here occurs at the base of the upper normal
subchron of magnetic Anomaly Sc (e.g., Chron CSCn), at about 16.5 Ma, in agreement
with the radiometric dates around this boundary (Hamor et al., 1987; Vass et al.,
1987). The base of the Serravallian stage was defined by Iaccarino (1985) at the base
of the Globorotalia praemenardi-Globorotalia peripheroronda subzone which corre-
sponds with the base of Blow plankton zone N10 and likewise correlates with the
lowermost normal of Chron CSA (e.g., CSADn).
The uppermost Badenian calcareous nannoplankton floras occur around and below
the NN6/NN7 zone boundary (Rogl and MUller, 1976). These biostratigraphic data
indicate an approximate position for the Badenian/Sarmatian stage boundary and are
supported by radiometric dates (Papp et al., 1974; Vass et al., 1987).
The middle/late Miocene boundary has been variously drawn in the last few
years. Published limits for this boundary range from 1Z.3 Ma up to 10.4 Ma, due to the
problems of correlation between magnetic Anomaly 5 and Chron 9 (resp. Chron 11)
which has been discussed above. The middle/late Miocene boundary is placed here at
·the base of the Tortonian Stage, within Blow plankton zone N15 (resp. NN9), and
coincides with the Chron 11/Chron 10 boundary; that is, the top of magnetic Anomaly
SA (equals CS/CSA). This gives an age of 11.5 Ma for this boundary.
The Sarmatian/Pannonian stage boundary in central European endemic sequences

can be estimated only by the FAD of Hipparion primigenium at the base of the
Pannonian stage (Bemor et al., 1988) and by radiometric dates (e.g., Hamor et al.,
1987; Papp et al., 1985; Vass et al., 1987). Therefore, this boundary is drawn here at
11.5 Ma.
The Pannonian/Pontian stage boundary is well defined lithostratigraphically,

biostratigraphically, and chronostratigraphically within the Central Paratethys (see
Papp et al., 1985; Steininger et al., 1985; Bemor et al., 1988a). However, a reliable
correlation of the Pannonian/Pontian stage boundary and the Pontian Stage with
Mediterranean stages is currently unresolved and quite controversial (see Steininger
and Papp, 1979; Rogl and Steininger, 1983; Bemor, 1983; Stevanovic, 1987; Bernor et
al., 1988 for a review of this problem). There are several MN11-13 mammal faunas
(see Steininger and Papp, 1979; Bernor et al., 1988) known from Pontian sediments in
Austria and Hungary. These mammal faunas, as well as a series of radiometric dates,
provide an estimate of the chronologie range of Pontian stage as about 8.0 to 5.8 Ma
(Semenenko, pers. comm., April Z8, 1988; Vass et al., 1987). A number of recent
paleomagnetic investigations, however, have yielded considerable interlaboratory
discrepancies and are therefore not included in this interpretation (Andreescu, 1981,
1987; Andreescu et al., 1987; Chepalyga et al., 1985; Pevzner, 1987; Pevzner and
Vangenheim, 1985a,b).
The base of the Messinian stage is drawn here at the base of the Globorotalia
conomiozea zone as defined by Iaccarino (1985), coinciding with the NNlla/NNllb
boundary as shown also by Berggren et al. (1985a,b), Barron et al. (1985a,b), and
Martini and Muller (1986). This boundary correlates with the base of the lower normal
subchron in Chron C3A, with an age of 6.5 Ma.
The Miocene/Pliocene boundary is drawn worldwide in terms of planktonic

biozonations at the N17/N18 (e.g., the NNll b/NN1Z) zone boundary which correlates
19
approximately with the Chron 5 (e.g., C3A) and Gilbert (e.g., C3) boundary, at
5.3 Ma. A chronostratigraphic boundary-stratotype is currently under discussion.
However, recent magnetostratigraphic studies of Zijderveld et al. (1986) have shown
that, in terms of Mediterranean stages, the Messinian/Zanclean stage boundary should
be placed just below the Thvera normal subchron (in the Gilbert chron) at 4.83 or
4.84 Ma. Channell et al. (1988) suggest an age of about 4.84 to 4.93 for the base of
the Zanclean stage. This does not necessarily imply the worldwide adjustment of the
accepted Miocene/Pliocene Series boundary with the Mediterranean Messinian/
Zanclean stage boundary (F .F. Steininger, pers. opinion).
The Zanclean/Piacencian stage boundary follows Iaccarino (1985) and Rio et al.
(1988).
The Pontian/Dacian (as well as Dacian/Romanian) stage boundaries follow

Bernor et al. (1988) and Andreescu (1981).
CONTINENTAL BIOZONA'llONS
Pollen Zonations
The regional pollen zonations shown here are directly correlated with marine
planktonic zonations covering most of the Neogene in the eastern Mediterranean
(Benda and Meulenkamp, 1979, 1989), the Pliocene-Pleistocene interval in the western
Mediterranean (Sue, 1982., 1987; Sue and Zagwijn, 1983), and the Oligocene and lower
Miocene in the central Paratethys Realm (Hochuli, 1978; Rogl et al., 1981). Direct
continental-marine correlations are provided through these pollen zonations and they
can be applied for a more precise correlation of mammal zonations with the marine
chronologie framework.
Mammal Zonations
The currently used European mammal biozonation follows recommendations

provided by international symposia (Munich 1975 and Madrid 1976) and the Neogene
Congresses of the Regional Committee on Mediterranean Neogene Stratigraphy held in
Bratislava 1975, Athens 1979, and Budapest 1985 (Alberdi and Aguirre, 1977;
Fahlbusch, 1976; Mein, 1975, 1981). The stratigraphic subdivisions used here are the
result of the Workshop on "European Neogene Mammal Chronology" held at Schloss
Reisensburg in 1988. At this meeting a lively discussion first cleared stratigraphic
concepts and the decision was taken to provide not only a biostratigraphic zonation,
but also an initial set of continental chronostratigraphic stages for scientific testing
(see below). A revised version of the currently used European mammal biozonation is
provided by P. Mein in this volume. In Mein's contribution the biologic criteria that
define and limit these zones are given from zone MN1 at the Oligocene/Miocene
boundary through MN17 at the Pliocene/Pleistocene boundary.
Neogene Mammal Fauaal Zones
These MN zones are named here by an informal biostratigraphic term: "Neogene

mammal faunal zones." The criteria defining these Neogene mammal faunal zones, as
given by P. Mein in this volume, are (1) the evolutionary FADs of taxa (mostly genera),
(2.) the FADs of immigrant taxa (mostly genera), (3) the LADs of selected taxa (mostly
genera), and (4) relatively common taxa within the zone. A reference locality con-
taining the characteristic mammal fauna of the zone and correlative localities located
throughout the circum-Mediterranean area are listed. Within the corresponding zone
these localities are listed (1) according to their geographic area and (2.) from bottom
to top of the zone according to their biochronologic position within the zone.
Neogene Mammal Farmal Units
Several of these "Neogene mammal faunal zones" have been combined into
20
larger units and a specific name has been given for some of these units (Fahlbusch,
1976). Since these units have been proposed entirely under biochronologic considera-
tions they are named here by an informal biochronologic term: "Neogene mammal
faunal units." To distinguish these "Neogene mammal faunal units" from the proposed
European continental chronostratigraphic stages and to avoid possible confusion, we
recognize the value of a proposal by M. Freudenthal (in letter dated December 31,
1988) to use instead of the ending -ian Hum, -ien), which is applied to chronostrati-
graphic terms, the ending -ic for these units. Following this proposal, the correct
writing and extent of these "Neogene mammal faunal units" should be: Agenic (MN1
and MNZ); Orleanic (MN3 to MN5); Astaracic (MN6 to MN8); Vallesic (MN9 and MN10);
Turolic (MNll to MN13); Ruscinic (MN14 to MN15). For the upper Pliocene, respec-
tively the faunal zones MN16 and MN17, the terms Villafranchic (commonly used in
western and southern Europe) and Villanyic (commonly used in central and eastern
Europe) are widely accepted, but neither was recognized by the Reisensburg
symposium. However, these sorts of nomenclature changes should be introduced only
after thorough discussion by a broad group of paleontologists and biostratigraphers.
There were several other proposals at the Reisensburg meeting for naming these
units: J. Agusti (Sabadell, Spain) would prefer "mammal unit" and "mammal super-
unit;" M. Freudenthal (Leiden, Netherlands) would prefer "mammal unit" and "mammal
episodes;" L.J. Flynn (Cambridge, USA) would prefer "Mammal Neogene Zones (or
Biochrons)" and "Mammal Ages;" E. Lindsay (Tucson, USA) "Mammal Neogene zones
(Mein, 1979)" and "Mammal Ages (Fahlbusch, 1975)." However, several other Neogene
mammalian zonations were proposed earlier and are still partly used: Aguilar (198Za),
Aguilar and Michaux (1984), Unay and de Bruijn (1984), and Thaler (1966). Revised
"Neogene mammal faunal zones" based on a rodent taxon range zonation for the upper
Miocene, Pliocene, and Pleistocene is proposed by Fejfar and Heinrich in this volume.
Earlier and more recent marine/continental-mammalian biochronology correla-

tions have been attempted by Aguilar (198Za), Aguilar and Michaux (1984), Azzarolli
et al. (1988), Berggren et al. (1985b), Cicha et al. (1972), Lopez-Martinez et al. (1987),
Papp (1981), Rogl and Steininger (1983), Steininger (1977), Steininger et al. (1985), and
Torre (1987).
The correlation presented here of the revised Neogene mammal faunal zones
(Mein, this volume; Fejfar and Heinrich, this volume) is based on a set of mammal
faunal localities, which provided more or less direct correlations to the marine bio-
chronologies, the magnetic time scale, the pollen zonations, and/or are located rela-
tive to radiometric dates. Only localities with significant mammal faunas have been
selected and their biochronologic position within the specific "Neogene mammal
faunal zone" was taken into consideration (see also Mein, this volume) •. The localities,
detailed criteria, and the literature used for these correlations are listed in another
section: "Data base for correlation tie-points."
Continental/Marine Correlations
Ageuian (MN1 and MNZ). The correlation of the lower boundary of zone MN1
cannot be solved for the moment and the exact position of the MN1 and MNZa zone
boundary in terms of marine correlation is also questionable. The MNZa/MNZb zone
boundary might coincide with the type Aquitanian. The localities Gans and Aillas
(lower MNZa) within the Aquitanian in the type area and Caunelles (higher MNZa)
within NN1 and the locality Balizac (lowermost MNZb) were recovered from marine
type Aquitanian sediments of zone NN1.
Orleanian (MN3 to MNS). Several localities (Estrepouy, Lisboa-Univ. Catolica

[I,II = R1], and Maigen) indicate the close correlation of the base of MN3 zone with the
base of the Burdigalian stage (as well as Eggenburgian). The lengthy duration of MN3
zone is supported by the locality Beaulieu (higher MN3) with radiometric dates 18.7
and 17.5 Ma. The locality Lis boa IVb (= RZ) (higher MN3) with marine correlations of
upper NN3 and NN4 zones and Blow plankton zone N7 to N8 would even indicate a
younger age for the top of MN3 zone. However, characteristic early MN4 zone local-
21
and 17.5 Ma. The locality Lis boa IVb (= R2) (higher MN3) with marine correlations of
upper NN3 and NN4 zones and Blow plankton zone N7 to N8 would even indicate a
younger age for the top of MN3 zone. However, characteristic early MN4 zone local-
ities (Quinta do Narigao, Lisboa Va (= R3), Aliveri-Kymi and Belenyenice) indicate a
correlation with NN3 to NN4 zones and Blow plankton zone N7 by marine biostratig-
raphy, pollen zonation, and radiometric dates. Furthermore, the FAD of European
Proboscidea can be biostratigraphically dated within NN3-zone (R:ogl and Steininger,
1?,83; Baldi, pers. comm.). The top of MN4 zone is well correlated by the locality
Orechov (middle to late MN4) coming from uppermost Ottnangian littoral marine
sediments. Also, Ziegler and Fahlbusch (1986) report three localities from the
Bavarian Molasse, Germany (Rauscherod, Rembach, and Forsthart) of later MN4 age
which are found stratigraphically higher than the Ottnangian Oncophora beds, and
hence would be correlated with the early Karpatian stage. This would imply that the
MN4/MN5 boundary would correlate within the Karpatian, respectively the late
Burdigalian marine stage (Fahlbusch and Bernor, pers. opinion). The base of zone MN5
is well correlated below the FAD of Globigerinoides bisphaericus (= base of Blow
plankton zone N8) at the locality Eibiswald (base of MN5) and, paleomagnetically, at
the locality Gemerek (base of MN5). A new locality Teiritzberg (middle to late MN5,
pers. comm. P. Mein and 0, Fejfar) was found in littoral marine to paralic sediments
of Karpatian Age. According to the localities like Quinta do Pombeiro (latest MN5),
Chelas 1 (late MN5), and St. Genies (probable late MN5), uppermost MN5 zone should
occur within the base of Blow plankton zone N8. According to the opinion of Bernor
(after Rogl and Steininger, 1983; Haq et al., 1987; Bernor et al., 1988b), MN 5 extends
to the late Langhian regression, respectively basal Middle Badenian, ca. 15 Ma;
accordingly, the base of MN6 would then be ca. 15 Ma. Some other authors report
potentially younger MN5 localities, including: Chelas 2 (within Blow plankton zone
N9/N10); Veyran correlated with nanno-zone NN6; Dumlupinar within the upper
Eskihisar pollen zone, and Chios (latest MN5 zone according to Mein; MN7 zone and
lower Yeni-Eskihisar pollen zone according to Benda and Meulenkamp, 1989).
The short duration of zones MN4 and MN5 might correspond to eustatically
controlled African-Eurasian mammal exchange, continuing into basal MN6 zone.
These faunal exchanges were interrupted by the basal Langhian/Badenian transgression
and followed by the late Langhian regression which permitted basal MN6 interconti-
nental mammal exchange (Bernor, 1983; Bernor et al., 1987; Rogl and Steininger, 1983;
Steininger et al., 1985a; Thomas, 1985). Bernor (pers. opinion) believes that current
evidence suggests that the base of MN5 zone correlates well with the terminal
Burdigalian regression (e.g., the Karpatian stage, about 16.5 Ma), and the base of the
MN6 correlates with the terminal Langhian regression, (resp. lower/middle Badenian
stage, about 15 Ma).
Astaracian (MN6 to MN8). The.fissure fillings of Neudorf Spalte 1-3 (Devinska

Nova Ves) have caused an endless discussion during the last decade and only some of
the latest opinions will be given here. Mein (1975) placed the Neudorf Spalte mammal
fauna at the base of his zone MN6. He moved it down to MN5 in 1981 (Mein, 1981) and
in this volume he placed the fauna back at the base of zone MN6, together with the
Turkish locality Pasalar. The fauna was recovered from several karstic fissure fillings
which are transgressively overlain by a marine, mica-rich, sandy to marly formation.
This marine formation is of middle to late Badenian age according to Cicha et al.
(1972). However, there are two marine transgressive events recognized on top of the
fissures according to Fejfar (Paratethys Symposium at Rohanov, CSSR, April 29, 1988
and Reisensburg meeting abstract). The first transgressive event correlates with the
lower Badenian stage (Lagenid zone) and the second transgressive event correlates
with the upper Badenian stage (Bulimina-Bolivina zone). Reevaluation of the state-
ments given by Fejfar, based on personal communication from Cicha (pers. comm.,
July 14, 1989), shows that only late Badenian marine sediments overlie the Neudorf
Spalte fissure fillings. Based on paleogeographic reasoning (Rogl and Steininger,
1983), the fissures were accessible only at the upper Burdigalian (upper Karpatian)
regressive event, or at the upper Langhian (= middle Badenian) regression. Following
the opinion of Bernor (above) and the evidence given at Neudorf Sandberg locality, we
place the base of MN6 within the middle Badenian stage, at approximately 15 Ma.
22
Other rather early MN6 faunas would also suggest this age and correlation; for
example, Luc sur Orbieu fauna (= Blow plankton zone N9/N1) and Pasalar fauna (=
lower Eskihisar pollen zone). The Neudorf Sandberg (= sandhill) locality also repre-
sents a rather early MN6 faunal interval. The mammal remains are found in marine
littoral sediments of middle to upper Badenian stage. The localities Steinberg and
Goldberg (middle MN6) appear to be younger than 14.7 Ma. ~andir (late MN6) shows
an Eskihisar pollen zone. The MN6/MN7 zone boundary occurs within Blow plankton
zone N11 according to the locality Santarem (latest MN6 or earliest MN7) and reaches
younger into Blow plankton zone N1Z (La Grenatiere =middle MN7; La Grive M =late
MN7). The locality Plakia (middle MN7) with early Yeni-Eskihisar pollen zone reflects
the problem of the MN7/MN8 zone boundary correlation to the marine sequence.
Early MN8 zone faunas include C. Almirall (NN7, especially N13/N14), Sankt Stefan
i.L. (lower Sarmatian (Volhynian) stage), Comanesti 1 (lower Sarmatian (Volhynian)
stage), Sof~a (Yeni Eskihisar pollen zone). The Yeni-Eskihisar 1 fauna (basal Yeni-
Eskihisar pollen zone) radiometric date (13.Z Ma) and Basarakavak fauna (upper Yeni-
Eskihisar pollen zone) and radiometric date (11.7 Ma) are limiting for younger MN8
zone correlations.
Vallesian (MN9 and MNlO). The FAD of Hipparion (= Cormohipparion, a junior

synonym of Hippotherium) was reevaluated by S. Sen (Reisensburg meeting) as no older
than 11.5 Ma. This is in accordance with early MN9 localities like Howenegg (radio-
metric dates between 1Z.4 and 10.8, most reliable date for Howenegg beds 10.8);
Esme-Akcakoy (radiometric date 11.6 ± 0.5 Ma), and Bou Hanifia (FAD of first local
Hipparion 100m above a radiometric date of 1Z.03 ± O.Z5 Ma). The Bou Hanifia
Hipparion site was estimated about 10.5 Ma by Sen, based on paleomagnetic data, and
the horse was considered derived based on morphologic data (by Bernor, Tobien and
Woodburne, this volume). From the Central Paratethys, lower Pannonian strata yield
radiometric dates of 11.6, 11.00, 10.7 Ma; middle to upper Pannonian strata yield
dates of 9.6, 8.4, 8.1 (Vass et al., 1987). The correlation of the Pannonian A to E (as
well as F to H) local zonation is discussed by Bernor et al. (1988). The fauna of the
basal MN9 locality Hovorany is placed with the Pannonian B/C local zone, the younger
MN9 locality Gaiselberg is placed within Pannonian C local zone. The MN9/MN10
zone boundary correlation with marine strata is established by the Kastellios (Greece)
localities with the lower part of Blow plankton zone N16, and with Chron C5r, just
before magnetic Anomaly 5, with the base of the Kizilhisar pollen zone. The mammal
remains of the localities Vosendorf and Inzersdorf (MN10 faunas) come from
Pannonian D toE local zone clays. Oued Zra (middle to late MN10) should be younger
than 9.7 ± 0.5 Ma. Levkon 1 (late MN10) is characterized by the lower Kizilhisar
pollen zone.
Turolian (MNll to MN13). The marine correlation of the base of the Turolian
mammal faunal unit can be estimated by the following localities: Crevilliente 1-3
(basal MN11) by Blow plankton zonation equals higher N16/N17 boundary; Kayadibi
(earliest MN11) equals lower Kizilhisar pollen zone with mammal fauna between radio-
metric dates of 9.4 and 7.95 Ma; Lower Maragheh (earliest MN11) is underlain by a
volcanic tuff dated 9.3 Ma; La Celia (= Los Gargantones) is overlain by lava beds with
a radiometric age of 7.Z to 7.6 Ma; Middle Maragheh (late MN11) position is between
volcanic tuffs dating 9.•3 and 7.75 Ma; Quarry X at Samos (early MN11) dated radio-
metrically at 8.5 Ma; Garkin (late MN11) by lower Kizilhisar pollen zone and radio-
metric date of 8.6 Ma below mammal-bearing beds. Correlation tie-points for
mammal faunal zone MN1Z are scarce. Casa del Acero (basal MN1Z zone) is on top of
the third evaporitic cycle (first evaporitic cycle occurs on top of marine clays of
Globorotalia conomiozea zone); Upper Maragheh (early MNlZ) associated with vol-
canics dating from 7.Z to 6.9 Ma. Kinik (middle MN1Z zone) is placed in the upper
Kizilihsar pollen zone along with the questionable relation of Crevillente 4 to the
plankton Globorotalia conomiozea zone. MN13 localities (Mein, this volume) ordered
biochronologically include the following controversial correlation tie-points. Molina
de Segura (earliest mammal fauna of MN13 zone) is on top of the third evaporitic
cycle, as above. Pikermi (MN1Z/MN13 boundary) is within the upper Kizilihisar pollen
zone; Samos (= Samos 5) (earliest MN13) occurs in a reversed polarity interval accord-
ing to Sen and Valet (1986) between 6.1-6.4 Ma; Venta del Moro (early to middle
23
MN13) is in Chron 5 (equals C3A) in normal part of base or top of magnetic Anomaly
3A; Crevillente 6 (middle MN13) is at the base of Messinian Stage with Globorotalia
conomiozea, and La Alberca (late MN13) is correlated with upper Messinian Stage.
Convincing evidence for a direct correlation of MN13 with the uppermost Messinian
Colombaci Formation, overlain by marine sediment yielding fossils of the
Sphaeroidinellopsis zone, were demonstrated at Monticino, Italy by Giuli and Vai
(1988).
Ruscinian (MN14 and MN15). Levels E to L of the Molina de Segura sequence,

which represent early MN14 faunas, are superposed on the Messinian evaporitic cycles;
all other MN14 faunas with possibilities for chronologie resolution represent middle
MN14 faunas. These include Baccinello V3 (overlain by Globorotalia margaritae-
bearing sediments); Elbistan (with lower Akca pollen zone and radiometric date of
3. 7 Ma); Dinar-Akcakoy, Karaburun, Spilia 1, and Ptolemais 1 (all with lower Akca
pollen zone); Terrats (placed in Gauss magnetic chron by Sue and Zagwijn (1983), but
within Gilbert magnetic chron by Opdyke and Lindsay (pers. comm., 1989) with Pll
pollen zone); Vendargues (base of Pll pollen zone). The MN14 and MN15 zone boundary
should occur near the Gauss (normal) and Gilbert (reversed) magnetic chron boundary
according to Opdyke and Lindsay (pers. comm., 1989). The localities Terrats (MN14),
Serrat d'en Vacquer (= Perpignan = MN15), and Sete (late MN15) are all characterized
by the Pll pollen zone.
'"Villafranchian• - '"Villanyian• (MN16 and MN17). The "Triversa faunas" (=

Villafranchian stratotype) of Italy (equal MN16) correlate best with the lower
Matuyama magnetic chron according to Opdyke and Lindsay (pers. comm., 1989). The
locality Arcille occurs above the Globorotalia punticulata zone; Vialette is underlain
by ash beds dated 3.3 to 2.6 Ma, it is therefore younger than 2.6 Ma; Fornace ROB
quarry has indirect correlation to the Globorotalia puncticulata/Gr. aemiliana (= Gr.
crassaformis) transition. Middle to late ("Montopoli-type faunas") of MN16 correla-
tions are Giilyazi (middle MN16), within the middle Akca pollen zone; Rincon 1 (late
MN16); Les Etouaires with radiometric dates of 2.6 to 2.4 Ma. Montopoli occurs above
the Globorotalia aemiliana (= Gr. crassaformis) zone within the lower Matuyama
magnetic chron. This indicatesthe MN16/MN17 zone boundary should occur in the
Matuyama magnetic chron, and the basal MN17 zone mammal faunas support this
correlation. For example, Villaroya with Pill pollen zone and lower Globorotalia
inflata zone occurs in reversed strata correlated with the Matuyama chron. Roca
Neyra has radiometric dates of 2.5-2.4 Ma and Saltina 2 is from reversed strata corre-
lated with the Matuyama magnetic chron. The best correlation for the MN16/MN 17
zone boundary is found in the Stranzendorf loess section. The mammal-bearing
horizons Stranzendorf A to C (= faunal zone MN16) are in strata with normal polarity,
possibly correlative with the Olduvai subchron; the horizon Stranzendorf D (already
faunal zone MN 17) was recovered from immediately over lying strata with reversed
polarity. The top of mammal faunal zone MN17 can be constrained in terms of radio-
metric dates: Chillac (late MN17) is underlain by basalts dated 1.9 Ma with reversed
polarity, as well as pollen zonations: Tegelen = Tiglian C (latest MN17) is correlated
with PIV - Pll pollen zones.
European Continental Chronostratigraphy
Chronostratigraphy subdivides Earth history, recorded in rocks and fossils, into

specific intervals of geologic time based entirely on rock sequences. Chronostrati-
graphic units are therefore bound, and characterized only by specifically choosen
sections and boundary stratotype points in the field. Each chronostratigraphic unit, as
a defined interval of rock strata, has a corresponding geochronologic unit, which
accounts for this specific interval of geologic time. A different chronostratigraphic-
geochronologic hierarchical system of terms is used to differentiate between these
two stratigraphic systems. The chronostratigraphic term "stage" and the geochrono-
logic term "age" serve as the basic units of these hierarchical systems (Hedberg,
1976).
24
In European as well as North American mammalian zonations the inconsistent
usage and mixture between geochronologic-chronostratigr aphic, biostratigraphic, and
biochronologic terms is still existant. This problem was addressed at the Reisensburg
meeting, and our correlation table (figure 1) is our attempt to clarify the distinction
between these stratigraphic terms. The extensive continental Neogene basins in Spain
(Steininger et al., 1985b) triggered and provided possibilities to develop chronostrati-
graphic stages for continental deposits in Europe. At the moment these stages are in
use only for the Iberian Peninsula. They should be tested for wider application, and
correlated possibly throughout the rest of Europe by mammal biochronology, and in
some cases by magnetostratigraphy. Chronostratigraphic mammal stages based on
Spanish sections have been proposed and their use is encouraged here to test their
application and utility in resolving paleontologic and geologic problems.
However, Neogene chronostratigraphic stages are, in general, confined region-

ally and mostly reflect the geologic history of a particular area. Only Neogene Series
and Subseries, the next higher ranking chronostratigraphic units, have a worldwide
application.
For the upper Oligocene, and probably extending into lower Miocene, no official
proposal has been made for a European mammal stage. However, Agusti et al. (1988,
and letter dated September Z7, 1988) are working on possible stratotype sections in
the Ebro Basin.
The Ramblian stage and its type section was established by Daams et al. (1987).
The Aragonian stage and its type section was published by Daams et al. (1977) and
further discussed by Daams and Freudenthal (1981, 1987). The Vallesian stage was
originally designated by Crusafont Pairo (1950) and redescribed by Marks (1971),
Aguirre et al. (1975), and Agusti (1988). Work on potential boundary stratotype
sections is in progress (Moya-Sola and Agusti, 1987). The Turolian stage was originally
designated by Crusafont Pairo (1965) and redescribed by Marks (1971) and Aguirre et
al. (1975).
Several participants at the Reisensburg meeting recommended that Vallesian and

Turolian stages be restudied and updated. Evaluation of the stratigraphic extension of
the Vallesian is especially in debate. J. Agusti favors the present biochronological
limits of the Vallesian including MN9 and MN10 faunal zones (letter dated September
Z7, 1988). S. Moya-Sola, emphasizing that the most important mammalian faunal
change in Spain takes place between faunal zone MN9 and MN 10, suggested two possi-
bilities: (1) to eliminate the Vallesian faunal unit, by moving MN9 faunal zone into the
Astaracian faunal unit, thereby extending the Aragonian superstage upward, and to
move MN10 faunal zone into the Turolian faunal unit thereby extending the Turolian
stage downward; or (Z) to conserve the Vallesian in terms of a valid faunal unit and
stage by restricting it to faunal zone MN9 (letter dated September ZZ, 1988). M.
Freudenthal proposed incorporating the lower Vallesian with the Aragonian and the
upper Vallesian with the Turolian (letter dated December 31, 1988). Bernor and
Fahlbusch comment that Freudenthal's and Moya-Sola's (number 1) proposals to
abandon recognition of the Vallesian faunal unit effectively eliminates one of the best
bases of intercontinental mammalian correlation (base of MN9) and subepoch boundary
definition (middle-late Miocene boundary closely corresponds) in the Neogene: the so-
called "Hipparion Datum." Bernor and Fahlbusch vigorously recommend the mainte-
nance of Vallesian and its contents, MN9 and MN10, according to the Munich
symposium, because of its chronological utility, and to avoid the unnecessary
confusion which would be caused by reassigning MN9 and MN10 to other MN units.
At present, the Reisensburg meeting does not recommend the official designa-
tion of any European Pliocene continental stages (e.g., Ruscinian: Kretzoi and Pecsi,
198Z; Villafranchian: Alberdi and Bonadonna, 1987; Kretzoi and Pecsi, 198Z;
Villanyian: Kretzoi and Pecsi, 198Z). This recommendation stems primarily from
absence of suitable stratigraphic sections where superposition of the rich and well
described Pliocene mammal faunas of Europe can be demonstrated.
25
ACKNOWLEDGMENTS
This work and accompanying chart represent a team effort; the contributions
and suggestions from all members of the team (mentioned below) are appreciated.
The project was initiated by Steininger at the suggestion of Fahlbusch. Steininger
prepared a preliminary chart and presented it at the Reisensburg meeting. The
preliminary chart was discussed vigorously, and the correlation committee (Steininger,
Flynn, Freudenthal, de Giuli, and Fejfar) continued that discussion, resulting in the
framework for the chart presented as table 1. Following the Reisensburg meeting,
Steininger sought input from additional colleagues, both Reisensburg meeting partici-
pants and other interested and experienced researchers. Notable additional sugges-
tions and revisions were submitted by Reisensburg participants Agusti, Benda, Bernor,
Freudenthal, Lindsay, Mein, Moya-Sola, and Sen, plus John Barron (U.S. Geological
Survey, Menlo Park, California, USA), Bill Berggren (Woods Hole Oceanographic
Institution, Massachusetts, USA), Gernot Rabeder (Institut fur Palaontologie, Univ.
Wien, Austria), and Fred Rogl (Naturhistorische Museum, Wien, Austria). The text was
prepared by Steininger, then scrutinized and polished through reviews by Bernor,
Fahlbusch, Mein, Lindsay, and Berggren. Typing was graciously provided by Ms. A.
Vogt, and drafting was skillfully provided by Mr. N. Frotzler.
As a team product, table 1 is compiled partly by concensus and surely must

include some errors. However, as a team product, it also represents current state-of-
the-art in chronological correlation, and we team members believe it represents the
most comprehensive, accurate, and current correlation of European Neogene chronol-
ogy. Its endurance will be a measure of its utility. We, as the preliminary compilers
of table 1, (along with the co-editors) are responsible for any omissions or errors that
were incorporated in the last hectic days of preparation. We sincerely thank all
members of the team mentioned above and all other participants of the Reisensburg
meeting who contributed their ideas and suggestions through discussions.
DATA BASE FOR CORRELATION TIE-POINTS
The mammal fauna localities in the following data base are arranged according
to their position in the mammal Neogene faunal zonation provided by P. Mein in this
volume. Stratigraphic positions according to other zonations and authors are given in
parentheses. The correlation tie-points for each locality follow the referenced
literature. The most recent literature available, especially summarizing papers, are
cited. However, since this data base list was collected and assembled by one person
(F .F. Steininger), many additional correlation points might be overlooked and others
not correctly interpreted. Improvement is implicit, and criticism is welcome. The
task has not been completed!
26
Table 1
Position
in Neogene
mammal faunal
zone accord-
Country, ing to Mein Correlation Literature etc. (latest
Locality (this volume) tie-points summarizing papers)
AGENIAN
France, MN1 -lower/ "Oligocene" Ringeade, 1978

Paulhiac middle
France, MNZa - lower- within transgression Hugueney and

Gans most of type Aquitanian Ringeade, 1988;
Ringeade, 1978
France, MNZa - lower- within transgression Hugueney and

Aillas most of type Aquitanian Ringeade, 1988;
Ringeade, 1978
France, MNZa- upper nanno-zonation NN 1 Aguilar 198Zb

Caunelles (Aguilar zone
A3)
France, MNZb - lower- marine type Alvinerie and Gayet,

Balizac most Aquitanian level 1971; Hugueney and
of Lariey Ringeade, 1988;
Ringeade, 1978
France, MNZb - middle marine type Hugueney and

Laugnac Aquitanian level Ringeade, 1988;
of Lariey Ringeade, 1978
France, MNZb marine type Hugueney and

La Brete Aquitanian level Ringeade, 1988;
of Lariey Ringeade, 1978
France, MNZb (Aquilar nanno-zonation NN 1 Aguilar 198Zb

Lespignan zone A4)
ORLEANIAN
Portugal, MN3 - lowermost on top of Aquitanian Antunes, 1988;

Lisboa Univ. (Aquilar zone transgression, over- Ginsburg, 1984;
Catolica AS) lain by marine Steininger et al.,
(I, n = R1) deposits; Blow 1987
zonation N4/N5;
nanno-zonation NNZ
Austria, MN3- lowermost nanno-zonation NNZ; Mein, 1989

Maigen Blow zonation N 5;
Central Paratethys
pollen zone NGZ n
Austria, MN3- basal Blow zonation NS; Hochuli, 1978; Daxner-

Eggenburg Central Paratethys Hock, 1971
(Brunnstube- pollen zone NGZ n
Schindergraben)
(cont.)
27
Table 1 (cont.)
Position
in Neogene
mammal faunal
zone accord-
France, MN3- basal above the type Gourinard et al., 1987;

Estrepouy Burdigalian trans- Hugueney and
gression (= "Marne Ringeade, 1988;
a Ostrea aginensis" Ringeade, 1978
with Miogypsina
globulina) in Peloua
marine level with
nanno zone NN2
France, MN3 (Aquilar radiometric dates: Aguilar, 1982b; Aguilar

Beaulieu zone B) 17.5, 18.7 Ma in Steininger et al.,
1987; Clauzon, 1982;
Clauzon and Aguilar,
1982
Portugal, MN3 Blow zonation N7/N8; Antunes, 1988; Aguilar

Lis boa IVb MN4 -lower- nanno-zonation NN4 in Steininger, 1987;
(=R2) most (Aguilar Ginsburg, 1984
zone B)
Greece, MN4 - lower to Kale-Eskihisar Benda and de Bruijn,

Aliveri-Kymi middle E-Med. pollen zone 1982; Benda and
Meulenkamp, 1989
Portugal, MN4- middle Blow zonation N8 Aguilar, 1982b;

Lisboa Va (Aguilar zone base; nanno- Ginsburg, 1984
(=R3) C1) zonation NN4 top
Czeshoslovakia, MN4- middle in littoral marine Cicha et al., 1972;

Orechov to upper facies below so- Fejfar, 1988
called "Oncophora
beds" = upper
Ottnangian
Turkey, MN4 (accord- lower Eskihisar Becker-Platen et al.,

Belenyenice ing to Benda E-Med. pollen zone; 1975; Benda and
and Meulenkamp, radiometric date at Meulenkamp, 1989;
1989) at mammal locality: Sickenberg et al.,
17.3 ± 0.4 Ma 1975
Turkey, MN 5 - lowermost according to de Bruijn et al., 1988

Gemerek magnetics, an
interpolated age
of 16.5 Ma
Austria, MN5- basal Karpatian Kollmann, 1965;

Eibiswald Rabeder and
Steininger, 1975
Austria, MN5- middle littoral marine to Grill, 1962; Sovis, 1987

Teiritzberg to upper (Mein, paralic Karpatian
pers. comm.)
28
Position
in Neogene
mammal faunal
zone accord-
Portugal, MN5- upper Chelas 1 - Blow Aguilar, 198Za; Aguilar

Chelas 1,Z (Aguilar: Chelas zonation N8/N9; in Steininger et al.,
1 = zone CZ; Chelas Z - Blow 1987
Chelas Z = zone zonation N9/N10;
C3) (equal to Blow zona-
tion N9 according to
Mein, pers. comm.,
1985)
Portugal, MN5- upper- Blow zonation N8 Aguilar in Steininger et

Quinta do most (Aguilar al., 1987; Ginsburg,
Pombeiro zone C1) 1984
France, MN5 top or MN6 nanno-zonation NN5 Aguilar, 198Za

Veyran base (Mein,
pers. comm.,
1989); Aquilar
zone CZ
Turkey, MN 5 (according upper Eskihisar Becker-Platen et al.,

Dumlupinar to Benda and E-Med. pollen zone; 197 5; Sickenberg et
Meulenkamp, radiometric date al., 1975; Benda and
1989) above mammal local- Meulenkamp, 1989
ity: 14.75 ± 0.3 Ma
Greece, MN5- upper lower Yeni-Eskihisar Benda and

Chios (according to E-Med. pollen zone Meulenkamp, 1989
Benda and
Meulenkamp,
1989; not MN 5
but MN7)
ASTARACIAN
Czechoslovakia, MN6- lowermost below lower Badenian Cicha et al., 197Z;

Neudorf- (Lagenid zone) Fej far, 1988; Rogl
Spalten 1-3 and Steininger, 1983
(Devinska
Nova Ves)
Turkey, MN6 - lowermost lowermost Eskihisar Benda and

Pasalar E-Med. pollen zone Meulenkamp, 1989;
Benda et al., 1975;
Sickenberg et al., 197 5
Czechoslovakia, MN6- basal middle to upper Cicha et al., 197Z;

Neudorf- Badenian Fejfar, 1988; Rabeder
Sandberg and Steininger, 1975
(Devinska
Nova Ves-
sandhill)
(cont.)
29
Table 1 (cont.)
Position
in Neogene
mammal faunal
zone accord-
Germany, MN6- middle younger than 14.7 Ma Heissig, 1988;

Steinberg and Heizmann and
Goldberg Fahlbusch, 1983
France, MN 6 (Aguilar Blow zonation Aguilar, 198Za

Luc sur zone C3) N9/N10
Orbieu
Turkey, MN 6 - uppermost Eskihisar E-Med. Benda and

)=andir pollen zone Meulemkamp, 1989;
Benda et al., 197 5;
Sickenberg et al., 19 7 5
France, MN7 - lower to Blow zonation N1Z; Aguilar, 198Za

La Grenatiere middle (Aquilar nanno-zonation NN 6
zone C4)
Greece, MN7 - lower to base of Yeni- Benda and

Plakia middle Eskihisar pollen Meulenkamp, 1989
zone
France, MN7 - middle to Blow zonation N1Z; Aguilar, 198Za

La Grive M upper (Aguilar nanno-zonation NN6
zone C4)
Portugal, MN7 (according Blow zonation N 11; Aguilar, 198Za

Santarem to Mein, pers. Nanno-zonation NN6
comm., 1989)
(Aguilar zone
C4)
Spain, MN 8 (Aguilar Blow zonation Aguilar, 198Za

C. Almirall zone C5) N13/N14 boundary;
Nanno-zonation NN7
Turkey, MN8 -lower Yeni-Eskihisar Benda and

Sofcta E-Med. pollen zone Meulenkamp, 1989;
Benda et al., 197 5;
Sickenberg et al., 197 5
Austria, MN8 -lower lower Sarmatian Mottl, 1975, 1980;

Sankt Stefan (= Volhynian) Rabeder and
i.L. Steininger, 1975
Romania, MN8 (according lower Sarmatian Feru et al., 1980;

Comanesti 1 to Feru et al., (= Volhynian, upper) Bernor et al., 1988a
1980)
30
Position
in Neogene
mammal falUlal
zone accord-
ColDltry, ing to Mein Correlation Literature etc. (latest
Turkey, Yeni- MN8 -uppermost Yeni-Eskihisar Becker-Platen et al.,

Eskihisar 1 E-Med. pollen zone; 197 5; Benda and
(locality radiometric dates: Meulenkamp, 1989;
number 2. with Yeni-Eskihisar 1 = Benda et al., 1975;
poor and 13.2. Ma; Yeni- Sickenberg et al.,
atypical Eskihisar 2. = 11.1 1975
mammal Ma
falUla)
VALLESIAN
Germany, MN9- basal several radiometric Zobelein, 1988; Bernor

Howenegg dates: 12..4 to 10.8 et al., 1988a
for basalts and
tuffs; most reliable
date for "Howenegg"
beds: 11.8 :1: 0.6 Ma
Turkey, MN9 (according radiometric date: Sen, 1988

Esme-Akcakoy to Fejfar, 11.6 ± 0.5
1989)
Algeria, MN 9- middle radiometric dates: Bernor et al., 1988a;

Bou Hanifia to upper 12..03 ± 0.2.5 Ma Sen, 1986
100m below lowest
Hipparions
Czechoslovakia, MN9 (basal Pannonian B/C local Bernor et al., 1988a;

Hovorany according to zone Ctyrocky, in Steininger
Bernor et al., et al., 1987
1988)
Romania, MN9- basal Pannonian C/D local Feru et al., 1980

Comanesti 2. (upper accord- z'one
ing to Bernor
et al., 1988)
Austria, MN9 - uppermost Pannonian C local Bernor et al., 1988a

Gaiselberg zone
Greece, MN9/MN10 Blow zonation N16; Benda and

Kastellios bolUldary (MN9 base of Kizilhisar Meulenkamp, 1989;
(2., 3) according to pollen zone; mag- Sen et al., 1986
Benda and netic assignment:
Meulenkamp, just below Anomaly
1989) 5 (Chron C5r)
Austria, MN10- basal Pannonian D toE Bernor et al., 1988a;

Vosendorf and local zones Rabeder, 1985;
Inzersdorf Rabeder and
Steininger, 1975
(cont.)
31
Table 1 (cont.)
Position
in Neogene
mammal faunal
zone accord-
Country, ing to Mein Cprrelation Literature etc. (latest
Morocco, MN10- middle radiometric date: Jaeger et al., 1973

Oued Zra to upper mammal locality
younger than
9.7 ± 0.5 Ma
Greece, MN10- upper lower Kizilhisar Benda and

Levkon 1 E-Med. pollen zone Meulenkamp, 1989
TUROLIAN
Spain, MN11- basal Blow zonation N16 Aguilar, 198Za

Crevillente (Aguilar zone
1-3 D3)
Turkey, MNll -lower- lower Kizilhisar Becker-Platen et al.,

Kayadibi most E-Med. pollen zone; 1975; Benda and
radiometric dates: Meulenkamp, 1989;
Bulumya ignimbrite Sickenberg et al.,
below mammal fauna: 1975
9.4 ± 0.2 Ma; Detse
ignimbrite above
mammal fauna:
7.95 ± 0.25 Ma
Iran, MNll -lower- mammal fauna under- Campbell et al., 1980;

Lower most lain by tuff dated Bernor, 1985, 1986
Maragheh 9.3 ± 0.1 Ma
Spain, MN 11 (accord- radiometric dates: Agusti in Steininger et

La Celia ing to Mein, lava beds above al., 1987
(=Los pers. comm., mammal locality:
Gargantones) 1989) 7.Z-7.6Ma
Turkey, MN11- upper lower Kizilhisar Becker-Platen et al.,

Gar kin E-Med. pollen zone; 1975; Benda and
radiometric dates: Meulenkamp, 1989;
8.6 Ma below mammal- Sickenberg et al.,
bearing beds 1975
Turkey, MNll- lower to middle to upper Benda and

Kiiciik- middle (MN1Z Kizilhisar E-Med. Meulenkamp, 1989;
Cekmece according to pollen zone Sickenberg et al., 197 5
Benda and
Meulenkamp,
1989)
Greece, MNll (accord- radiometric dates Salounias, 1981; Bernor

Samos (lower ing to Bernor near 8.5 Ma et al., 1980; Weidmann
fossil beds, et al., 1980) et al., 1984
e.g., Quarry
X)
32
Position
in Neogene
mammal fannal
zone accord-
Conntry, ing to Mein Correlation Literature etc•. (latest
Spain, MN1Z- basal over lies third Agusti in Steininger

Casa del Messinian evaporitic et al., 1987
Acero cycle (first
evaporitic cycle
overlies Globorotalia
conomiozea zone)
Turkey, MN1Z- middle upper Kizilhisar Benda and

Kinik E-Med. pollen zone Meulenkamp, 1989;
Sickenberg et al., 1975
Spain, MN13 -lower- overlies third Agusti in Steininger et

Molina de most (MN1Z/13 Messinian al., 1987
Segura bonndary evaporitic cycle
horizon D according to
and 1 Agusti, 1987)
Greece, MN1Z/MN13 uppermost Benda and

Pikermi bonndary Kizikhisar E-Med. Meulenkamp, 1989
pollen zone
Greece, MN13- basal within reversed Sen and Valet, 1986;

Samos (upper (MN 1Z accord- polarity zone, Weidmann et al., 1984
mammalian ing to Weidmann approximately
fossiliferous et al., 1984, 6.1 - 6.4 Ma
levels) and Bernor et
al., 1980)
Spain, MN13 -lower within a normal Opdyke et al., 1988

Venta del to middle polarity zone of
Moro Anomaly 3A
Spain, MN13 - middle lower Messinian, Aguilar, 198Za;

Crevillente (Aguilar zone Globorotalia de Bruijn et al., 197 5
6 EZ) conomiozea zone
Spain, MN13- middle mammal locality Aguil~,198Za;

Librilla to upper nnderlain by Montenat et al., 1975
(Aguilar zone volcanic nnit
EZ) dated 6.9 Ma
Spain, MN13 (upper Blow zonation N18; Aguilar, 198Za; Mein et

La Alberca MN13 according between marine sedi- al., 1973
to Fejfar, ments of Messinian
this volume; age; last occurrence
Aguilar zone of Discoaster
E3) guingueramus =
upper Messinian
(cont.)
33
Table 1 tcont)
Position
in Neogene
mammal faunal
zone accord-
Country, ing to Mein Correlation Literature etc. Oatest
Greece, MN13 (accord- uppermost Benda and

Rem a ing to Benda Kizilhisar E-Med. Meulenkamp, 1989
Marmara and Meulenkamp, pollen zone
1989)
Spain, MN13 (accord- over lies third Agusti in Steininger et

La Hornera ing to Agusti, evaporitic cycle; al., 1987
1987) above Globorotalia
conomiozea zone
Italy MN13 (accord- overlain by lower- Giuli and Vai, 1988

Monticino ing to Giuli most marine
quarry, et al., 1988) Zanclean (MPL 1,
Brisighella NNlZ, and Thvera
magnetic subchron
RUSCINIAN
Spain, MN13/MN14 over lies third Agusti in Steininger et

Molina de (according to Messinian al., 1987
Segura Agusti, 1987) evaporitic cycle; in
horizon E continuous section
to 10 with MN1Z/MN13
fauna
Italy, MN14 -middle overlain by marine Torre, 1987

Baccinello V3 (MN13/MN14 beds with
boundary accord- Globorotalia
ing to Torre, margaritae
1987; upper
MN13 according
to Fejfar and
Heinrich, this
volume)
Turkey, MN14 -lower lower Akca E-Med. Benda and

Dinar- pollen zone Meulenkamp, 1989;
Akcakoy Sickenberg et al., 1975
France, MN14 -lower NW-Med. pollen Aguilar and Michaux,

Terrats (MN14a accord- zonation PII zone; 1984; Sue and Zagwijn,
ing to Fejfar normal event in 1983; Lindsay, 1985
and Heinrich, Gilbert magnetic
this volume; chron
Aguilar and
Michaux, 1984,
zone FZ)
34
Position
in Neogene
mammal faunal
zone accord-
Greece, MN14- middle lower Akca E-Med. Benda and

Ptolemais 1 (MN14b accord- pollen zone Meulenkamp, 1989
ing to Fejfar
and Heinrich,
this volume)
France, MN14- middle NW-Med pollen Aguilar and Michaux,

Vendargues (MN14b accord- zonation base of 1984; Sue and Zagwijn,
ing to Fejfar Pn zone; reverse 1983; Lindsay, 1985
and Heinrich, polarity of Gilbert
this volume; magnetic chron
Aguilar and
Michaux zone
FZ)
Turkey, MN14 (accord- lower Akca E-Med. Benda and

Elbistan ing Benda and pollen zone; radio- Meulenkamp, 1989;
Meulenkamp, metric date: 3.7 Ma Sickenberg et al.,
1989) 1975
Greece, MN14 (accord- lower Akca E-Med. Benda and

Karaburun ing to Benda pollen zone Meulenkamp, 1989
and Meulenkamp,
1989)
Greece, MN14b (accord- lowermost Akca Benda and

Spilia 1 ing to Fejfar E-Med. pollen zone Meulenkamp, 1989
and Heinrich,
this volume)
France, MN15- middle NW-Med. pollen zona- Aguilar and Michaux,

Perpignan (Aguilar and tion Pn zone; under- 1984; Sue and Zagwijn,
(=Serrat Michaux zone lain by Gilbert 1983; Lindsay, 1985;
d'en Vacquer F3 to B1 magnetic chron
France, MN14 -lower NW-Med. pollen zona- Aguilar and Michaux,

Celleneuve (Aguilar and tion Pn zone; 1984; Sue, 1980, 198Z;
Michaux zone reversed polarity, Sue arid Zagwijn, 1983;
Gl) Gilbert magnetic Lindsay, 1985
chron
France, MN15- upper MN-Med. pollen Aguilar and Michaux,

Sete (Aguilar and zonation Pn zone 1984; Sue, 1980, 198Z;
Michaux zone G1) Sue and Zagwijn, 1983
Spain, MN14 base to lower Gauss Opdyke and Mein, 1988

Villalba alta MN15 top (Mein, magnetic chron
to Orios 3 pers. comm., (Anomaly ZA base)
(one section) 1989)
(cont.)
35
Table 1 (cont.):
Position
in Neogene
mammal faunal
zone accord-
"W.LAFRANCHIAN" - "W.LANYIAN"
France, MN16 -lower ash bed underlies Alberdi and Bonadonna,

Vialette mammal locality with 1987; Azzaroli et al.,
radiometric dates 1988; Bandet et al.,
3.3 to Z.6 Ma 1978; Ly Meng Hour et
al., 1983
Italy, MN16 -lower magnetic: reverse de Giuli et al., 1984;

Triversa polarity in Fornace Lindsay et al., 1980;
RDB quarry (= Torre, 1987
Matuyama magnetic
chron); indirectly:
Globorotalia
puncticulata/
Glr. crassaeformis
transition
Italy, MN16 -lower with Globorotalia Alberdi and Bonadonna,

Poggio aemiliana 1987
Mirteto (=crassaformis)
(Rom a) zone; ash radiometric
date: 3.3Z t 0.3;
reversed magnetic
polarity
Italy, MN16 -lower overlies marine Torre, 1987

Arcille sediments with
Globorotalia
puncticulata zone;
continental sediments
with mammal fauna
Tur~!!Y' MN16- middle middle Akca E-Med. Benda and

Gulyazi pollen zone Meulenkamp, 1989;
Sickenberg et al., 1975
Spain, MN16- middle magnetic normal Alberdi et al., 198Z;

Casa del polarity Azzaroli, 1988; Leone,
Rincon 1985; Torre, 1987
(Rincon 1)
France, MN16- upper ash below fossil bed Alberdi and Bonadonna,
Les Etouaires dated by several 1987; Azzaroli, 1988;
authors between 3.6 Lindsay et al., 1980
and Z.4 Ma; magnetics
in the area: Gauss
and Matuyama chrons
36
Position
in Neogene
mammal faunal
zone accord-
Italy, MN16- upper above marine sedi- Azzaroli, 1988; de Giuli

Montopoli ments with et al., 1984; Lindsay et
Globorotalia al., 1980; Torre, 1987
crassaformis; mag-
netic: at Gauss/
Matuyama boundary
= Z.45 Ma
Romania, MN16 (according magnetic: in lower Andreescu et al., 1981

Slatina 1 to Andreescu Matuyama magnetic
et al., 1981) chron; endemic
mollusc faunas =
Romanian Stage
Greece, MN16 (according lower part of Benda et al., 1977;

Rhodos to Benda et Globorotalia Benda and
(Kritka Fm.) al., 1977) inflata zone; nanno- Meulenkamp, 1989
zonation upper MN 16;
middle Akca E-Med.
pollen zone
Netherlands, MN16 (upper NW-Med. pollen Sue and Zagwijn, 1983

De Meeren MN16 according zonation: Pin/
(drill site) to Fejfar and PIV-Pl 1 zones
Heinrich, this
volume)
Austria, MN16 - upper- magnetic: normal Rabeder, 1981

Stranzendorf most (according event in Matuyama
horizon A-C to Rabeder, magnetic chron
pers. comm.,
1989)
Austria, MN17- from magnetic: reversed Rabeder, 1981

Stranzendorf lower to upper polarity of Matuyama
horizon D (according to magnetic chron
toM Rabeder, pers.
comm., 1989)
France, MN17 -lower- radiometric date: Azzaroli et al., 1988;

Roca Neyra most (MN16 Z.4 to z.s Ma Torre, 1987
according to
Azzaroli et
al., 1988)
Spain, MN17 -lower- NW-Med. pollen Sue and Zagwijn, 1983

Villaroya most zone zonation Pill
Romania, MN17 -lower magnetic: lower part Andreescu et al., 1981

Slatina Z of Matuyama chron
(cont.)
37
Table 1 (cone.)
Position
in Neogene
mammal faunal
zone accord-
Count,ry, ingto Mein Correlation Literature etc. (latest
France, MN17- upper underlain by basalt Azzaroli et al., 1988;

Chillac flows with reverse Boeuf, 1983
magnetization, dated
at 1.9 Ma
Austria, MN 17 - upper- magnetic: in Olduvai Rabeder, 1981

Stranzendorf most (according event of Matuyama
horizon M to Rabeder, chron
pers. comm.,
1989)
Nether lands, MN 17 - upper- NW-Med. pollen Sue and Zagwijn, 1983

Tiglian C most zonation PIV-Pl I
zone
38
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Thaler, L., 1966. Les rongeurs fossiles du Bas Languedoc dans leur rapports avec
l'histoire des faunes et la stratigraphie du Tertiaire d'Europe. Mem. Mus. Nat.
Hist. Nat. Paris, ser. C 17, p. 1-296.
Theyer, F. and Hammond, S.R., 1974. Paleomagnetic polarity sequence and radio-
larian zones, Brunhes to Polarity Epoch ZO. Earth and Planetary Science
Letters, v. ZZ, p. 307-319.
Theyer, F., Mato, C.Y., and Hammond, S.R., 1978. Paleomagnetic and geochronologic
calibration of latest Oligocene to Pliocene radiolarian events, equatorial
Pacific. Marine Micropaleontology, v. 3, p. 377-395.
Thomas, H., 1985. The lower and middle Miocene land connection of the Afro-Arabian
plate and Asia: A major event for hominoid dispersal?, in Delson, E. (ed.),
"Paleoanthropology: The Hard Evidence." Alan R. Liss, Inc., New York, p. 18.
Torre, D., 1987. Pliocene and Pleistocene marine-continental correlations. Ann. Inst.
Geol. Publ. Hung., v. LXX, p. 71-77.
Unay, E. and Bruijn, H. de, 1984. On some Neogene rodent assemblages from both
sides of the Dardanelles, Turkey. Newsl. Stratigr., v. 13(3), p. 119-13Z.
Vass, D., Repok, I., Balogh, K., and Halmai, J., 1987. Revised radiometric time-scale
for the Central Paratethyan Neogene. Ann. Inst. Geol. Publ. Hung., v. LXX, p.
4Z3-434.
Weidmann, M., Solounias, N., Drake, R.E., and Curtis, G.H., 1984. Neogene stratig-
raphy of the eastern basin, Samos Island, Greece. Geobios, v. 17 (4), p. 4 77-490.
Ziegler, R. and Fahlbusch, V., 1986. Kleinsauger-Faunen aus der basalen Oberen
Siisswasser-Molasse Niederbayerns. Zitteliana, v. 14, p. 3-58.
Zijderveld, J.D.A., Zachariasse, J.W., Verhallen, P.J.J.M., and Hilgen, F.J., 1986. The
age of the Miocene-Pliocene boundary. Newsl. Stratigr., v. 16(3), p. 169-181.
Zobelein, H.K., 1988. Die jungtertiaren Hoewenegg-Schichen im Hegau (Baden-
Wiirttemberg) und ihre Umgebung nach der Literatur. Mitt. Bayer. Staatsslg.
Palaont. hist. Geol., v. Z8, p. 173-186.
46
A BIOCHRONOLOGIC SUBDIVISON OF THE EUROPEAN
PALEOGENE BASED ON MAMMALS- REPORT ON RESULTS OF
THE PALEOGENE SYMPOSIUM HELD IN MAINZ IN FEBRUARY 1987
Norbert Scbmidt-Kittler
Institut fiir Geowissenschaften, Palaontologie

Johannes Gutenberg Universitat
Saarstr. Z1
Postfach 3980
D-6500 Mainz, FRG
INTRODUCTION
In February 1987 the paleontologists working on European mammalian faunas of

the Paleogene convened in Mainz to establish a mammal chronology for this time
period. The results were edited by the author of this report at the end of the same
year (Schmidt-Kittler, 1987). The biochronological subdivision agreed upon in this
symposium corresponds to the general conditions met in the European continental
Tertiary. Therefore, it is consistent with the concept of subdivision used for the
Neogene mammal chronology of Europe elaborated in an earlier meeting in Munich
(Fahlbusch, 1976).
The decision on a most appropriate and generally applicable time scale for the
European bioprovince based on mammals was made in clear view of the possibilities
and limitations found in the continental Paleogene of this region due to the particular
geologic conditions. These are, first, the existence of only few continental sedimen-
tary basins with a potential of mammal-bearing geologic sections, and, second, by far
the larger percentage of the rich mammal localities in Europe either cannot be
connected to well defined stratigraphic sections (isolated localities) or are preserved
without geological time indication (fissure fillings). Nevertheless, these deficiencies
in the documentation of European mammalian faunas did not hamper progress very
much in elaborating chronologies. This is due to the fact that many phylogenetic
lineages could be recognized in the European mammalian faunas. Thus, irreversible
evolutionary change could be used as another reliable criterion of time succession
independently from superposition of geologic strata.
Of course, it would have been preferable to base the European mammal chronol-
ogy on geologic sections. The advantage of such a system of subdivision is clear
because it allows the mammalian biochronological and paleoecological data to be
combined with results obtained from other groups of organisms, and also to be com-
pared with absolute time calibrations which may be possible in some sections in the
future. The main problem is that mammal localities in continental sedimentary
sections are normally scarce, irregularly distributed along the time axis and mostly
rather poor in fossil content. These natural attributes of most continental sections
are not so important in the event that many sites are available so that the data they
provide can be combined. This, for instance, is usually the case in the North Americaft
Tertiary where many temporal-overlapping continental sedimentary basins exist. In
Europe, however, the number of stratigraphic sections with good potential for

mammal faunas is too low and is restricted to only few regions. Moreover, the exist-
ing sequences do not cover the whole time span of the Paleogene. As a consequence,
besides remaining very incomplete, a mammal chronology based exclusively on geolog-
ical stages would be too coarse to provide a suitable basis for further studies of the
development of paleofaunas.
The choice of a biochronologic time scale for European Paleogene mammalian

faunas therefore is not the result of a preexisting concept but is simply the conse-
quence of what we have to work with.
And it must be explicitly stated here that the description of mammal-bearing

sections will be of great importance for our future work and should be established
wherever reasonably possible. The reason for this is that the development of
mammalian paleofaunas can be integrated to a more general view of the geologic,
organismic, paleoclimatic, and paleogeographic context only inasmuch as it will be
possible to connect mammalian data to other information preserved in bodies of rock
strata.
THE MAINZ SYMPOSIUM
In view of these necessities and possibilities the strategy agreed upon by the
members of the Mainz symposium was to establish a detailed generally applicable
mammal biochronology for the European Paleogene and then to add to it stratigraphic
sections where possible. On the grounds explained above, both procedures are equally
indispensible for further progress in studies of mammalian evolution. The first allows
precision in distance correlation; the second is a precondition for combining our
mammalian data with further information of the fossil record and the geologic back-
ground.
Table 1 displays the mammal biochronology elaborated by the participants of the

Mainz symposium. The designated reference levels are based on rich mammalian
faunas comprising both micromammals and macromammals. Though defined in this
form during the Mainz symposium, a first version of this subdivision already existed
for quite a long time. It had been worked out by a small group of Paleogene
30 CODERET
29 RICKENBACH
28 PECH DU FAAYSSE Chattian
27 BONINGEN
-------- r--- -----

28 MAS DE PAUFFIE
25 GAAOUILLAS OLIOGOCENE
24 HEIMERSHEIM
23 ITARDIES
22 VILLEBRAMAR Starn pian Rupelian
21 SOUMAILLE
GRANDE COUPURE
Priabonian
~
20 ST. CAPRAISE D. E.
19 ESCAMPS
(Ludian) Latdorfian
18 LA DEBRUGE
17 FONS4
16 ROBIAC
15 LA LIVINIERE Bartonian Bartonian s.l.
14 EGERKINGEN a + ~ EOCENE
13 GEISELTAL OMK (OBERE MITTELKOHLE)
12 GEISELTAL UMK (UNTERE MITTELKOHLE) Lutetian
11 GEISELTAL UK(UNTERKOHLE)
10 GAAUVES
8·9 AVENAY Ypresian
7 DORMAAL
6 CERNAY
1·5 HAININ PALEOCENE
Table 1. Biochronological subdivision of the European continental Paleogene

based on mammals as proposed by the International Symposium in
Mainz, February 1987. Sequence of faunal reference (MP) levels are
numbered to the left.
48
specialists during the 1975 Munich symposium (Fahlbusch, 1976). As an historical fact,
it must be said that chronological successions in the Paleogene of Europe were mainly
recognized at that , time using evolutionary stages of lineages. Extraction of
mammalian data from a sedimentary sequence was realized only on the Isle of Wight,
leading to the definition of the Headonian stage of the Oligocene (Bosma, 1974). Some
years before, however, the same time span had already been correctly subdivided in
other regions on the basis of rodent lineages described from fissure fillings (Schmidt-
Kittler, 1971; de Bonis et al., 1973; Hartenberger, 1973).
Since the elaboration of the first biochronologic approach in 1975 many new
important faunas were discovered and several mammal•bearing stratigraphic
sequences were described (Russell et al., 198Z; Engesser et al., 1984; Agusti et al.,
1985; Hooker, 1986; Alvarez-Sierra et al., 1987). And, in all cases where the existing
biochronology could be tested later by the geologic superposition of mammalian
faunas, it proved to be correct. In no case, however, did the biostratigraphic data go
beyond the standard of precision already attained by the mammal chronology using
only evolutionary information. This is not astonishing since chronologie successions
can be worked out more accurately when more faunas with statistically defined evolu-
tionary stages of species become known. And, in the European Paleogene many rich
faunas with statistically significant population sizes of rodents are available from
fissure fillings. Of course, events of immigration and extinction of micromammals
and macromammals are used for subdivision as well. These, however, can only be
taken as reliable inasmuch as they are controlled by the information drawn from
evolutionary lineages.
EUROPEAN PALEOGENE MAMMAL SEQUENCE
The faunas of the reference localities (table 1) form points on the time axis, not
intervals. Thus, no boundaries between the reference levels are defined and any
faunas added later to the succession by correlation must be assigned to a particular
level on the basis of their affinities as expressed by evolutionary stages, and first or
last occurrence of species. The fact that the intervals between the reference levels
are not defined does not diminish the accuracy of the system of subdivision because
this depends exclusively on the quality of the reference faunas.
Judging from the very detailed documentation of lineages, fossil information in

the European Paleogene is particularly rich for the periods MP 16-18 and MP 2.5·30.
On the other hand, due to the geological conditions in Europe, documentation of
mammalian faunas older than middle Eocene is much more scattered than in the
following period of the Paleogene and it is evident that the Paleocene actually corre-
sponds to a big gap in the mammalian report. This is why the numbers 0-5 were
reserved for sites which we hope will be discovered in the future. The levels MP 8 and
MP 9 are provisionally not separated because in the present state of research it is
difficult to choose the most significant among several successive faunas for a
European reference level.
A characteristic of the European bioprovince in the Paleogene is the marked

faunal endemism during long periods of the late Eocene and Oligocene. They were
partly caused by zoogeographical barriers (e.g., the Upper Rhine graben, at times
connected to the Molasse sea) and partly due to physical differences, such as lowlands
with fairly high ground-water levels (e.g., continental Molasse basins, Hampshire
basin, Paris basin) and dry karst highlands (Quercy, Suebian, and Franconian Alb).
Climatic gradients may have played an important role as well. The differences in
composition of the geographically restricted mammalian faunas caused by these
factors become even more pronounced through faunal documentation due to particular
geological settings in the European subprovinces. This is why regional biostratigraphic
frameworks are needed in addition to the more general biochronologic subdivision of
the European Paleogene mammal sequence.
49
REFERENCES
Agusti, J., Anadon, P., Arbiol, S., and Cabrera, L., 1985. Biozonacion mediante
Roedores (Mammalia) del transito Oligocene•Mioceno en el sector sureste de la
cuenca del Ebro. Paleont. i. Evol., v. 18, p. 131-149.
Alvarez Sierra, M.A., Daams, R., Lacomba, J.I., Lopez Marinez, N., Sacristan Martin,
M.A., 1987. Succession of micro mammal faunas in the Oligocene of Spain, in
Schmidt-Kittler, N. (ed.), "International Symposium on Mammalian Biostratig-
raphy and Paleoecology of the European Paleogene, Mainz, February 18-Z1,
1987." M\inchner Geowiss. Abh., (A), v. 10, 31Z p.
Bonis, L. de, Chrochet, J.-Y., Rage, J.-Cl, Sige, B., and Vianey-Liaud, M., 1973.
Nouvelles faunes de vertebres oligocenes des Phosphorites du Quercy. Bull. Mus.
natl. Hist. nat. Sci. Terre Z8, v. 174(3), P• 105-113.
Bosma, A.A., 1974. Rodent biostratigraphy of the Eocene-Oligocene transitional
strata of the Isle of Wight. Utrecht micropaleont. Bull., spec. Publ. 1, 1Z8 P•
Engesser, B., Mayo, N.A., and Weidmann, M., 1984. Nouveaux gisements de
mammeferes dans la Molasse subalpine vaudoise et fribourgeoise. Mem. suisses
Paleont. 107, 39 P•
Fahlbusch, V., 1976. Report on the International Symposium on Mammalian Stratig-
raphy of the European Tertiary. Newsletter Stratigraphy, v. S(Z/3), p. 16D-167.
Hartenberger, J.•L., 1973. Etude systematique des Tberidomyoidea (Rodentia) de
l'Eo~ene superieur. Mem. Soc. geol. Fr., (NS), 5Z (Mem. 117), 76 p.
Hooker, J.J., 1986. Mammals from the Bartonian (middle/late Eocene) of the
Hampshire Basin, southern England. Bull. Br. Mus. nat. Hist. (Geol)., v. 39, p.
191-478.
Russell, D.E., Hartenberger, J-L., Pomerol, c., Sen, s., Schmidt-Kittler, N., and
Vianey-Liaud, M., 198Z. Mammals and stratigraphy: The Paleogene of Europe.
Palaeovertebrata, Mem. extra, 77 p.
Schmidt-Kittler, N., 1971. Odontologische Untersuchungen an Pseudosciuriden
(Rodentia, Mammalia) des Alttertiars. Abh. Bayer. Akad. Wiss., math.-
naturwiss. Kl., N.F., 150, 133 p.
Schmidt-Kittler, N. (ed.), 1987. International Symposium on Mammalian Biostratig-
raphy and Paleoecology of the European Paleogene, Mainz, February 18·Z1,
1987. Miinchner Geowiss. Abh., (A), v. 10, 31Z p.
50
THE RAMBLIAN AND THE ARAGONIAN: LIMITS, SUBDMSION,
GEOGRAPffiCAL AND TEMPORAL EXTENSION
Remmert Daams
Museo Nacional de Ciencias Naturales

Jose Gutierrez Abascal 2
28006 Madrid, Spain
Matthijs Freudenthal
Rijksmuseum van Geologie en Mineralogie

Hooglandsehe Kerkgracht 17
2312 HS Leiden, The Netherlands
INTRODUCTION
In this paper a historical review of the definition and subdivison of the Aragonian
is given. Furthermore, the limits of the Ramblian and the Aragonian, and the
temporal and geographic extension of both stages are discussed. The "cricetid
vacuum" in the Lower Miocene of Spain is compared to that of France and central
Europe.
ffiSTORICAL REVIEW
The International Symposium on Mammalian Stratigraphy of the European

Tertiary, celebrated in April 1975 at Munich, decided that the Aragonian should
comprise beds between the entry of Anchitherium and that of Hipparion in Europe.
The type section was to be situated in the Calatayud-Teruel Basin. This proposal was
published by Fahlbusch (1976) as such (figure 1).
In 1977, Daams, Freudenthal, and Van de Weerd defined the Aragonian based on
collections made by de Bruijn and Freudenthal in the beginning of the sixties. The
stage was subdivided into a lower, a middle, and an upper portion.
In 1976, Daams and Freudenthal renewed excavations in the Calatayud-Teruel

Basin in order to obtain a more extensive documentation of the Aragonian in its type
area. A detailed subdivision was given by the same authors in 1981. It consists of a
lower part (Zone A), a middle part (Zones B, C, D, and E), and an upper portion (Zones
F and G). The Lower Vallesian was subdivided into Zones Hand I.
Daams and VanderMeulen (1983) created three pre-Aragonian zones (X, Y, and
Z) for fossiliferous sediments in the north of Spain. These zones were provisionally
assigned to the Agenian. At that moment only Zone X contained the fauna of Autol
(Ebro Basin, La Rioja), Zone Y that of Cetina de Aragon (Almazan Basin, Zaragoza),
and Zone Z the faunas from the Calamocha area in the Calatayud-Teruel Basin
(Navarrete; Ramblar 1, 3B, 4A, 5, 7, Valhondo 1).

Edited by E. H. Lindsay el at.
Daams
Daams
Freudenthal a Oaams a Daams a Freudenthal a Daams a
Fahlbusch Freudenthal Van der Meulen Alvarez Freudentha I
Van de Weerd
1976 1981 1983 1987 8
r:r 1977
_j _j _j ...
I-
I I I
..J
~
..J
~ H
..J
~
j -
H
..J
~
I 1--
..5! H
z ~
ci z ... .8: z ... G
8: -
.. z
8:-
..... G
<(
.
z ...
8:r-
G
<( en <( <( <(
_...::;_
~
- r- -
~
F
- r- -F
~
- ~
F
z - -
z z<( z z E z E .. - E
0 z 0 .!! 0 .. -
:0 D
0
-
..
:0 D
0
(!)
:0
·e
'0
D
- -
'0
·e ·e -c
(!) (!) (!)
r-
..
'0
(!) <( '0 :5! <(
I.LI <( <( <( E c
a:: ... c
•-
<(
modern ..J a:: r- a:: r- <(
B B ..5! B
c ricetids. h a:: a:: <( ~ r- <( ---.: r-
<( 0 ..5!1 A .21 A ai o."" A
~~ , ... -
Anchitheriu111
z .z z a:: .5!1 z
r-
Ritteneria
<(
-z
<(
I
y
z
I.LI I.LI
(!)
-
(!)
X
Rhodanomys <(
....._ ~
Fig. 1. Chronological history of the definition of limits and subdivision of the

Ramblian, Aragonian, and Lower Vallesian.
The presence of many pre-Aragonian faunas in the Calamocha area demanded

the creation of a new stage. As the Agenian was never properly defined as a stage, it
was decided to define the Ramblian stage for a sequence which was originally meant
to be approximately the Late Agenian. The Ramblian Stage (Daams et al., 1987) was
defined to comprise the sequence and faunas recorded after the extinction level of
Ritteneria to the entry of the first modern cricetid (Democricetodon). The Ramblian
consists of a lower zone, and an upper one containing the "cricetid-vacuum." In the
same paper, the lower limit of Aragonian Stage was redefined as the entry of
Democricetodon. The Lower Aragonian would contain Zones B and C (as determined
by Daams and Freudenthal, 1981), and the Middle Aragonian Zones D and E. The
further subdivision of the Upper Aragonian and Lower Vallesian was not changed.
Daams and Freudenthal (1988) maintain the same limits and main subdivisions,
but divide both Zones D and G into three subzones.
THE SUBDIVISION OF THE RAMBLIAN
The Ramblian contains Zones Z and A. Zone Z is characterized by the absence

of Ritteneria, by the initiation of the radiation of Ligerimys (Alvarez, 1987), by a
diverse fauna of Gliridae, and by the presence of Eucricetodon aff. aquitanicus and E.
aff. infralactorensis. The lowermost fauna of this zone (Navarrete) contains abundant
Eucricetodon, and the uppermost faunas have few specimens of this genus. A diverse
record of Sciuridae (5 species) is characteristic for this zone as well (Cuenca, 1988).
Melissiodon may be represented in low numbers.
The following zone (A) is characterized by a diverse Eomyidae record (48-84% of

the rodent fauna). The Gliridae and Sciuridae are both qualitatively and quantitatively
somewhat impoverished, but Glirudinus modestus is relatively well represented.
Melissiodon may be present in low numbers, and in the lowermost fauna (La Dehesa)
one tooth of Eucricetodon is present. The other, generally more diverse, faunas are
52
void of Eucricetodon, and consequently represent the "cricetid-vacuum." The lower
limit of this zone is not a sharp one, the fauna of La Dehesa being transitional. The
presence of Eucricetodon and of only one species of Ligerimys shows affinities with
Zone z, but the large quantity of Eomyidae teeth {48%) and the quantitatively
impoverished Gliridae association show more affinity with Zone A.
For further details on the subdivision of the Ramblian, the location of its type
area, and its type section, the reader is referred to Daams et al. {1987).
THE RAMBLIAN/ARAGONIAN LIMIT
Daams and Freudenthal {1981) and Daams et al. {1987) mentioned the rare occur-
rence of Anchitherium in the Aragonian type area and adjacent areas. As a matter of
fact, only two remains of this equid have been found during the 1976-83 excavations.
One was found at an isolated place in the type section, and the other find is from
Moratilla {Zone A). For us this was reason enough to abandon Anchitherium as a
criterion for defining a stage limit. In the type area of the Ramblian Anchitherium is
not present among the macromammal remains in faunas from Zone Z {Morales and
Soria, 1984). The above mentioned considerations made us abandon the entry of this
equid as a criterion for the Ramblian/Aragonian boundary {Daams et al., 1987). The
stage limit was moved upward and is now placed at the entry of Democricetodon, the
first modem cricetid in central and western Europe.
SUBDIVISIONS OF THE ARAGONIAN AND THE LOWER VALLESIA.N
The subdivision of the Aragonian and Lower Vallesian has been exhaustively
dealt with in Daams and Freudenthal (1981), Daams et al. {1987), and Daams and
Freudenthal {1988). The reader is directed to those papers for details of this subdivi-
sion. A few of these details are repeated here for clarification. Zones D and G have
each been subdivided into three subzones by Daams and Freudenthal {1988). Subzone
D1 contains abundant Fahlbuschia koenigswaldi, in subzone DZ Pseudofahlbuschia
jordensi is well represented, and D3 is characterized by Fahlbuschia freudenthali and
Renzimys lacombai. Zone G is subdivided into three subzones on the basis of the
evolutionary stages of the large-sized Megacricetodon lineage. Subzone G 1 houses M.
collongensis-crusafonti, GZ contains M. crusafonti, and G3 is characterized by M.
crusafonti-ibericus.
THE ARAGONIAN/VALLESIAN LIMIT
This boundary was defined previously by the supposed simultaneous entry of

Hipparion into the Old World. We now think that this has been an unfortunate
choice. First of all, it has never been proven that the Hipparion event is synchronous;
it has only been an assumption. Second, micromammal remains are much more abun-
dant than large mammal ones and provide a more robust biostratigraphic framework.
J;..opez et al. {1986) showed that the first occurrence of Hipparion in the faunal succes-
sions of various Spanish basins {figure Z) is diachronous, and consequently does not
serve as a criterion for a stage limit. Therefore, the Aragonian/Vallesian boundary
should be reconsidered. We would recommend it be established on the basis of one or
more micromammal events, recognizable in the entire Old World, or at least Europe.
This event may be present below or above the actual "boundary," and it is stressed
that it should be defined in a section where both Upper Aragonian and Lower Vallesian
faunas are present in stratigraphic superposition. These circumstances are not
currently represented in the Calatayud-Teruel Basin, nor in Catalonia, but they are
favorable in the Duero Basin. Unfortunately, the faunas in the Duero Basin are not as
rich as they are in the Calatayud-Teruel Basin, but those from the Torremormojon and
Ampudia sections {see L6pez-Martlnez et al., 1986 and Gm:da Moreno, 1987) are
sufficiently represented to define in more detail the faunal succession around the
Aragonian/Vallesian boundary. One could argue that Spain is in a non-central geo-
53
(11
BIOSTRATIGRAPHY BIOCHRONOLOGY
"'" CASTILLA ARAGON I CATALUNYA SPAIN TURKEY
I I
I
Progonomys Progonomys Progonomys Progonomys (Progonomys and
hispanic us hispanicus 1 hispanic us hispanicus Parapodemus)
Biozone Biozone Biozone Biocronozone Biocronozone E
I
(Progonomys)
Biocronozone nlf-
1--------1--- -
'
Cricetulodon Biozone Cricetulodon Cricetulodon (Cricetulodon)

hartenbergeri hartenbergeri
I Biozone
Biozone • Biocronozone Bioc.,..onozone C
I
---- ---
Megacricetodon Biozone Fahlbuschia Megacricetodon (Megacricetodon)
ibericus I crusafonti
I H ibericus
I
Biocronozone B
Biozone I Biozone • Biocronozone
• - - - - - - -•
Megacricetodon Biozone Faunas with two
lopezae species of
Biozone G
Megacricetodon
Biozone
F
I
Alvarez et al. Daams & Freuden- Agust.!, 1981 L6pez et al.,l986 Unay & De Bruijn,
1986 thal, 1981; Van 1984
de Weerd, 1976
Fig. 2.. Biostratigraphy and biochronology of the Aragonian/Vallesian transition in Spain and Turkey. The star reflects
the first fossil record of Hipparion (slightly emended after Lopez et al., 1986).
graphical position for establishing a type zone for chronostratigraphic units. On the
other hand, there are few, if any, alternatives in Europe. In Spain, the conditions of
exposure are excellent, and there are many sections where faunas are unquestionably
superposed.
In this paper we do not want to change the Aragonian/Vallesian boundary, nor do

we have a proposal for an alternative. But we emphasize the need for a reconsidera-
tion of this limit.
DISCUSSION OF THE •cRICETID VACUUM•
In the fairly complete succession of Ramblian faunas in the Calamocha area, it

appears that Eucricetodon gradually decreases in number to disappear at the base of
Zone A. This decrease coincides with an increase of Eomyidae. Expansion of
Eomyidae was interpreted by Van de Weerd and Daams (1978) and Daams and Van der
Meulen (1984) by expansion of a more forested biotope, and consequently E. aff.
aquitanicus and~· aff~ infralactorensis may have preferred dry habitats. This hypoth-
esis is supported by the abundance of ~· aguitanicus in the fissure filling of Bouzigues,
fissure fillings representing a relatively dry biotope (Van de Weerd and Daams, 1978).
Generally MN Zone 3 is considered to represent the "cricetid vacuum." In

France, the locality of Estrepouy is placed in this zone, in spite of recording abundant
E. infralactorensis. Daams et al. (1987) mentioned the correlational problems between
MN zones and the succession in the Spanish Neogene. The fauna of Estrepouy may be
correlated to Zone z, because of general resemblances. Other French faunas corre-
lated to MN Zone 3 are those of Petit Camon, Richevoltes, Mounicot, Marsolan, and
Navere 1 (Baudelot and Collier, 1978, 1982.). Most of these localities contain low
numbers of~· infralactorensis, with the exception of Richevoltes where this species is
absent. Eomyidae are also present, but according to Baudelot and Collier (1978) this
family is predominant in Mounicot only. These faunas may be placed in either Zone Z
or A. Unfortunately, the quantitative and qualitative data on these faunas are not
complete,.and a more precise correlation is not possible.
Aguilar et al. (1986) mention new localities in the eastern French Pyrenees, of
which Sainte Catherine z, 3, 4, 5, and 6 are void of Eucricetodon and modern
cricetids. These localities may be correlative with our Zone z. In Sainte Catherine 7
only one modern cricetid (Democricetodon) is present, and hence this fauna may be
correlative with our Zone B. Serre de Verges (Meurisse et al., 1969) is another locality
belonging to the "cricetid vacuum."
In Catalonia, Agustf (1981) places four faunas in the Pseudodryomys ibericus

zone, which he in turn correlates with MN Zone 3b. These faunas are Sant Andreu de
la Barca 1 and 3b, Moll Calopa and Les Forques. · However, each of these contains only
a few teeth which are not considered to be representative. Therefore, the Catalonian
succession is not discussed here.
In Portugal, several lower Miocene faunas are known as well. But the only one
probably belonging to the "cricetid vacuum" contains only a few teeth, and is there-
fore not discussed here.
In Germany, the fissure fillings of Bissingen, Wintershof-West, and Schnaitheim

with abundant Eomyidae are void of Eucricetodon and modem cricetids as well.
Daams et al. (1987) correlated these faunas with Zone A. The Petersbuch Z and
Erkertshofen faunas (Ziegler and Fahlbusch, 1986) record Democricetodon as the only
modern cricetid. These two faunas may be correlative with our Zone B.
In Czechoslovakia, the locality of Tucho;ice is void of Eucricetodon and it has

abundant Eomyidae (Fejfar, 1974); it may therefore be correlative with our Zone A.
The succeeding faunas of Dolnice 1 and Z house Democricetodon as the only modern
cricetid, and are consequently correlative with our Zone B.
55
U1
en
CENTRAL SPAIN FRANCE' GERMANY CZ E CHOSLOVAK IA
' l.'ILLAFELICHE. 2.A 1...

-~-·
:.~-; ...
~s-~N ·Roou.E 2'.:/-
B
;~;~;~~~:·~{~;/~ ·.. t .JA!tf;~::.:. · ERKERTSHOFEN : ~ DOLNICE 2.
:eLMO REDON DO 3
. ·:;_t..' 7--~: ·, . ·. ·.. '
I.PETERSBUCH 2 ;;;,.- ,BbL~-~~~--. 1 · .-.:.~~J OEMO CR.
t~~~;f&~fG.-i~}. ·_. ·.-~ . rtr:~:~:~:;~:,: ·.<-: • • • • • ·- . • •
1 I
.....----:--
.. . • • • • • • • •• • 0 • • •
:l.l q~A;T.II,.I,.~ :: ~ ~ ~ ~
:-:-> -:.;-:-: sAINTE;.;-:-:·
• • • • • r • •' •' • • ..
"~-~-~-~~i~~i~·M·: ::::: ~ ..... ..
..·.·." .. . ....-.-.-:
............ ..
·BANON 11:-:-: -:-:-;, -:.:-CATH ER INE:.;.;. : ;:;: TUCHO ICE ;:;::"CRICETID"
::::: 2., 3,4,5,6 :::::: l:~~~~~~s~?~_:_~~s:T: · ••••.·.· •••••· •·••••.••· • VACUUM
A :BISS I NGEN · ::::::::::: ~
'sER ~E ·DE. VERGE·s·

.·.·.·.·.·.·.· ... ..... ... I
.. . ......... ..... ..·.·.·.·.·. .:
~
I
~ ~ ~
~t~~\~~;)} ~;:} I. • • • • ~ •' • • • • • , -··.. • .• • • • • • • • • !:::::::::::::::::;:::::::: :: ~
" . .
...
.. .................. ..... ........ .
~
.• ••·.• • ~ ~
· AGREDA ·. ·.. ,·.·.
.. . . .. . .
·LA DE HESA -:.:. NAVERE 1 :-:. EUCRI C.
'RAMBLAR 5 .... MARSOLAN ....

. .
. .... ;, . .. . 7 . . . . . . ' MOUNI CO l .. . _
.. .. · .. · .. .. . 38' . .. . · R ICH EVOLTE S :-
. ·. · , ;, . ·. · . '4A. ·. · . PE T IT CAMON ·
z ·VALHONoo · 3A. · . ·
'· •.. ESTREPOUY . ...
:. : . ~ ··! . : . :. : 1 • . : • :
-~A !'I~ L :4~ 1 : . :- :- ? r.:- :.:- :.:- :.:.· ..
'NAVARR-ETE • ·• · . · • - -·-- . - . · . ·. L AUGNAC L _ __ _ _j
Fig. 3. Faunal successions, in which the "cricetid-vacuum" is present, in various European countries.
Figure 3 summarizes the faunal sequence in the various European areas. In
central Spain, the succession seems fairly complete, although we do not know where
we should place faunas such as Ateca 1, 3 and Rubielos de Mora. Anyway, the gradual
decrease of Eucricetodon, followed by the predominance of Emyidae, and the subse-
quent entry of Democricetodon is clear in central Spain. In the south of France, the
gradual decrease of Eucricetodon seems apparent also, but quantitative and qualita-
tive data are insufficient to verify if Eomyidae are subsequently the abundant, and
diverse family. Only one fauna with one modern cricetid is present. Faunas in
Germany and Czechoslovakia with abundant Eomyidae and absence of Eucricetodon
are followed by those containing only one modern cricetid.
It is not possible to say if the "cricetid vacuum" is of the same duration in all
these mentioned areas, because many of these faunas lack detailed quantitative and
qualitative analysis. In Spain, the "cricetid vacuum" has the duration of more or less
one zone (A), but in terms of the MN zonation it is of half a zone (3b). We do not feel
the data are currently adequate to express these durations in millions of years.
GEOGRAPHIC EXTENSION OF THE RAMBLIAN AND THE ARAGONIAN
It is stressed here that these stages are primarily defined as chronostratigrahic

units. It is implicit that a strong to partial similarity of faunal elements should be
present, if these terms are to be applied outside of Spain. If not so, Ramblian and
Aragonian should not be applied. Within Europe, there are sufficient resemblances
among contemporaneous faunas, at the generic or specific levels, to correlate to the
type Ramblian and type Aragonian. Another objection is that when correlating over
such large distances, the error may increase considerably.
Comparing the micromammal faunas from both sides of the Dardanelles (Unay
and de Bruijn, 1984) of Turkey to the central and western European record, it appears
that faunal differences prevail over faunal similarities. These authors part from the
assumption that Hipparion arrived simultaneously throughout the Old World. However,
Lopez et al. (1986) show that the first fossil record of Hipparion, even within Spain,
may be diachronous. The micromammal faunas from Turkey have several genera and
species in common with western and central European faunas. However, these genera
and species are not adequate to serve as a solid basis for direct correlation. A sound
basis for application of the term Aragonian to faunal associations without Hipparion is
not present in Turkey. This basis is more solid for the Vallesian faunas, as there is
more general resemblance between the Spanish and the Turkish faunas of this age.
Concluding, it may be said that with the present state of knowledge the terms
Ramblian and Aragonian should only be applied for mammal chronology in Europe.
THE TEMPORAL EXTENSION OF THE RAMBLIAN AND THE ARAGONIAN
The temporal extension of these stages must be extracted indirectly from other
sources. Correlation tables such as the one in Rogl and Steininger (1983) include
Mein's (197 5, 1979) zonation as the main reference for continental events. If their
correlations are more or less correct, the base of the Ramblian is set at ZZ.5 Ma, and
its top at 18.5 Ma, implying a duration of 4 million years for this stage. The top of the
Aragonian would be situated at 10.5-11 Ma, and therefore the Aragonian would have a
duration of 7.5-8 million years. The Lower, the Middle, and the Upper Aragonian
would have a duration of 1, Z, and 4.5-5 million years, respectively.
However, we doubt these broad correlations. Daams and Freudenthal (1981) and
Daams et al. (1987) mentioned the limited usefulness of Mein's (1975) zonation on a
European scale. Also, the error increases in correlating from a radiometric scale
through MN zonations to local faunal successions. A more precise biostratigraphic
correlation should be established among the various European bioprovinces, using both
qualitative and quantitative data, before a more exact limit of the duration of these
stages can be obtained.
67
ACKNOWLEDGMENTS
We thank Dr. Nieves Lopez-Martinez (Madrid) for critically reading the manu-
script and for valuable suggestions. The results obtained from the Spanish succession
find their origin in excavations carried out from 1976-1986, which were financed by
the "Rijksmuseum van Geologie en Mineralogie" at Leiden (The Netherlands), the State
University of Groningen (The Netherlands), The Netherlands Organization for the
Advancement of Pure Research (Z.W.O.), and the "Instituto Geologico y Minero de
Espaiia (I.G.M.E.)."
REFERENCES
Aguilar, J.-P., Calvet, M., and Michaux, J., 1986. Decouvertes de faunes de
micromammiferes dans les Pyrenees-Orientales (France) de !'Oligocene superieur
au Miocene superieur; especes nouvelles et reflexion sur l'etallonage des echelles
continentale et marine. C. R. Acad. Sc., 303, 11(8), p. 755-760.
Agustr, J ., 1981. Roedores Miomorfos del N eogeno de Cataluna. Tesis Doctoral,
Barcelona, Z93 p.
Alvarez Sierra, M.A., 1987. Estudio sistematico y bioestratigrafico de los Eomyidae
(Rodentia, Mammalia) del Oligoceno superior y Mioceno inferior espanol.
Scripta Geologica, 86, Z07 p.
Baudelot, S. and Collier, A., 1978. Les faunes miocenes du Haut Armagnac (Gers,
France). 1. Les gisements. Bull. Soc. d'Hist. Nat. Toulouse, v. 114(1-Z), p.
194-Z06.
Baudelot, S. and Collier, A., 198Z. Les faunes de mammiferes mio.cEmes du Haut-
Armagnac (Gers, France): Les Glirides (Mammalia, Rodentia). Geobios, v. 15(5),
p. 705-7Z7.
Bruijn, H. de and Moltzer, J.G., 1974. The rodents from Rubielos de Mora; the first
evidence of the existence of different biotopes in the Early Miocene of eastern
Spain. Proc. Kon. Ned. Akad. Wet., B, 77(Z), p. 1Z9-145.
Cuenca Bescos, G., 1985. Estudio paleontologico de los Roedores (Mammalia) del
Mioceno inferior de Autol (La Rioja). Estudios Riojanos, Z, 96 p.
Cuenca Besc6s, G., 1988. Los Sciuridos del Aragoniense y del Rambliense en la Fosa
de Calatayud-Montalban. Scripta Geologica, 87, 116 p.
mammal zones. Scripta Geologica, 6Z, 17 p.
Daams, R. and Freudenthal, M., 1988. Synopsis of the Dutch-Spanish collaboration
program in the Aragonian type area. Script Geologica, Special Issue 1, p. 3-18.
Daams, R. and Meulen, A.J. van der, 1983. Paleoecological interpretation of micro-
mammal faunal successions in the Upper Oligocene and Miocene of Spain.
Mediterranean Neogene Continental Paleoenvironments and Paleoclimatic
Evolution, R.C.M.N .S. Interim Coll., Montpellier, April 1983, 4 p.
Daams, R. and Meulen, A.J. van der, 1984. Paleoenvironmental and paleoclimatic
interpretation of micromammal faunal successions in the Upper Oligocene and
Miocene of north central Spain. PaH!obiologie Continentale, v. 14(Z), p. Z41-Z57.
Daams, R., Freudenthal, M., and Weerd, A. van de, 1977. Aragonian, a new stage for
continental deposits of Miocene age. Newsl. Stratigr., v. 6(1), p. 4Z-55.
Daams, R., Freudenthal, M., and Alvarez Sierra, M.A., 1987. Ramblian; a new stage
for continental deposits of early Miocene age. Geologie en Mijnbouw, v. 65, p.
Z97-308.
Fahlbusch, V., 1966. Cricetidae (Rodentia, Mammalia) aus der mittelmiociinen
Spaltenfiillung Erkertshofen bei Eichsfatt. Mitt. Bayer. Staatsslg. Palaont. hist.
Geol., v. 6, p. 109-131.
raphy of the European Tertiary. Newsl. Stratigr., v. 5, p. 160-167.
Fejfar, 0., 1974. Die Eomyiden und Cricetiden (Rodentia, Mammalia) des Mioziins der
Tschechoslowakei. Palaeontographica, A, v. 146, p. 100-180.
Garda Moreno, E., 1987. Roedores y Lagomorfos del Mioceno de la Zona central de la
Cuenca del Duero. Sisteqratica, Bioestratigrafia y Paleoecologfa. Ph.D. Thesis
(inedited), Universidad Complutense, Madrid, Z58 p.
58
~opez, N., Garcfa, E., and Alvarez, M.A., 1986. Paleontol0 gia y Bioestratigraffa
(micromamlferos) del Mioceno medio y superior del sector central de la Cuerca
del Duero. Studia Geol. Salmant., v. ZZ, p. 191-Z1Z.
Mein, P., 1975. Resultats du Groupe de Travail des Vertebres. Report on Activity of
the R.C.M.N.S. Working Groups, Bratislava, p. 77-81.
a
Mein, P., 1979. Rapport d'activite du groupe de travail vertebres. Mise jour de la
biosratigraphie du Neogene basee sur les mammiferes. Ann. Geol. Pays Hellen.,
H.S., 7th Intern. Congr. R.C.M.N.S., Athens, 1979, v. 3, p. 1367-137Z.
Meurisse, M., Michaux, J ., and Sige, B., 1969. Un remplissage karstique a
micromammiferes du Miocene inferieur a la Serre de Verges, pres St.-Arnac
(Pyrenees-orientales). c. R. Soc. Geol. Fr., v. 5, p. 166-168.
Morales, J. and Soria, D., 1984. Los Artiod4ctilos del Mioceno inferior de las Cuencas
centrales de Espana. COL-PA, v. 39, p. 51-59.
Rogl, F. and Steininger, F., 1983. Vom Zerfall der Tethys zu Mediterran und
Paratethys. Die Neogene Pafaogeographie und Palinspastik des zirkum-
mediterranen Raum. Ann. Naturhist. Mus. Wien, v. 85/A, p. 135-163.
sides of the Dardanelles, Turkey. Newsl. Stratigr., v. 13(3), p. 119-13Z.
Weerd, A. van de and Daams, R., 1978. Quantitative composition of rodent faunas in
the Spanish Neogene and paleoecological implications. Proc. Kon. Ned. Akad.
Wet., B., v. 81(4),p. 448-473.
Ziegler, R. and Fahlbusch, V., 1986. Kleinsauger Faunen aus der basalen Oberen
Sii8.$wasser-Molasse Niederbayerns. Zitteliana, v. 14, p. 3-80.
59
NEW NEOGENE RODENT ASSEMBLAGES FROM ANATOUA (TURKEY)
Muhsin SfuQengen1 , Engin Unay1 , Gercek Sarac1 ,

Hans de Bruijn2 , Ismail Terlemez1 , and Mustafa GUrb1iz1
lM.T .A. Genel MiidiirlUgii

Ankara, Turkey
2Instituut voor Aardwetenschappen

3508 TA-Utrecht
The Netherlands
INTRODUCTION
Mapping projects in continental Tertiary basins under the auspices of the M.T.A.
(Mineral Exploration and Research Institute of Turkey) led in recent years to the
discovery of many sites yielding small mammals from formations that were previously
considered to be unfossiliferous. This study gives a brief account of the lithostratig-
raphy and biostratigraphy of the continental deposits of the southern part of the
Kayseri-Sivas basin. The rodent assemblages collected in that area are ranging in age
from the Middle Oligocene to the early Pliocene, showing a gap in the early Miocene.
We have filled this hiatus in our knowledge by studying two assemblages from succes-
sive levels in the lignite quarry of Harami (Akljehir area).
The qualitative and quantitative composition of our rodent associations will be

given and the potential of the Anatolian succession for establishing biostratigraphical
correlations with the Standard MN-zonation and with some Asiatic sequences will be
discussed.
THE GEOLOGY OF THE GEMEREK AREA
The area studied is part of the Kayseri-Sivas basin (figure 1). This NE-SW trend-
ing basin is delimited by the metamorphic rocks of the Akdag in the north and of the
Hinzirdag in the south.
In the southern part of the study area, south of the Elmah Dag thrust (figure 1),
the metamorphics are overlain by a series of turbiditic deposits of Eocene age (Erhan,
M.T.A. internal report) known as the Malakoy Formation. North of the Elmah Dag
thrust the basement is overlain by the pyroclastics, olistostromes and sandstones of
the yaldag group.
The Malakoy Formation is overlain by the Tuzhisar member of the Ceviz~ik

Formation, a sequence consisting of gypsum, sandstones and siltstones. The gypsum
deposits that seem to document the regressive phase of the marine Eocene are quite
thick southwest of Ortakoy, but wedge out toward the northeast. Upward the gypsum
deposit grades into the cyclic clastic fluviatile sediments of the D"okmetafi member of
t~~ Ceviz<;ik Formation (figure 2). The minimum thickness measured for the
Dokmetafi member is 500 m.

Edited by E.H. Lindsay et a/.
Ol
N
Horomi
lignite q110ay
..: [Upper [B) limestone

·~ U . Miocene
:3" Lower~ C:~91omerot,., sand.stone,
L::J a11tstooe and c:loystone
..: Mid. Miocene
~ rUpper[[l] ~~~~:·~~~um
-~ Lower~ sandstone, sttW,on~.morl
~ limestone and n.gnite
,..i
~[Low. -Mld. Miocene
i' QYplum, siltstone .,.
5 ~ and sandstone ·[~~ U.Oiigo-M.Miocene?
m ~ jj ~~ conglomerate. tondslone
I.&.. ~ EL.::!....l and cJoyatone
"2 , . U.Eocene- Oiigocene?
i j t [±J gypsum, siltttone
..l(- Anticline
Eocene u ~E and sandstone
~ Syncline
C'-4o~~ok\y9';:,-::. ~ =~~~~~u:li~~=m~!rts, tubKSites thrust fault
\7\7 \7 \l
rodent localities \7\7'\7, + +
Basement metamorphic rocks
g
....• + +
Fig. 1. Geological map of the Gemet;~k area (Kayseri;Sivgs basin). The rodent localities are indic~ted by
the numbers 1-8. 1 = Yenikoy; Z = Horlak 1 , 1 ; 3 = Horlak Z; 4 = Gemerek; 5 = Kalekoy; 6 =
Karaozii; 7 = Dendil; 8 = Igdeli.
Upper Oligocene_Middle Miocene?
3 ~~-------------------------------i
;.ID:;~ I Dokmeta~ member .._

~~ f. South of Elmal1 Dag t;rust fault.
f oJ Lower _Middle g
~ Miocene Middle Miocene ::> Upper Miocene Pliocene
:.
~
at
~
~----------------------~~----~
~, g Burtepe Form. Yenicubuk Formation J I Egerci Formation w
I I lower I upper. lower !upper_ North of Elmal1 Dag
N
f ff ff f t .. - thrust fault.
0
0 :I: COo
? 0 ~g' ~~ ~ c.
~ ~3 mm ~ <1>
Q) ~ "0'
~ o=""
"' "m
1\)"' '< C::
-
"'
CT
-
Fig. z. Diagram giving the relative positions of the formations and the rodent localities in the Gemerek area. All the age
assignments of continental deposits are based on fossil rodents.
0)
Col
In the northern part of the study area (north of the Elmah Dag thrust fault) the
lithology of the basin infill is rather different. The oldest Neogene unit recognized in
this region is the Burtepe Formation which crops out in the central part of the
Yeni~ubuk anticline. Lithologically the Burtepe Formation resembles the Tuzhisar
member of the Ceviz~ik Formation although sandstones seem to be less common.
Finds of small mammals (see below) show conclusively that the Burtepe Formation is
of Middle Miocene age whereas the Tuzhisar member is of Oligocene age.
The Burtepe Formation is conformably overlain by a sequence of sedimentary

cycles consisting of sandstone, marl, lignite and limestone: the lower Yenifubuk
Formation. The upper Yeniyubuk Formation consists of alternating siltstone, lime-
stone and gypsum beds. The biotite from the two basalt flows occurring in this unit
above the rodent localities Gemerek and Horlak Z (figure 1) were dated in the Z.W.O.
laboratory for isotope geology (Amsterdam) at 14.9 :1: 0.7 Ma and at 16.0 :1: 0.5 Ma,
respectively.
The Yeni~ubuk Formation is unconformably overlain by the conglomerates and

sandstones of the Egerci Formation. The lower part of this unit is coarse-grained and
at least 500 m thick in the northeastern part of the area, but much finer grained and
thinner in the southwestern part. Evidently the source area of these clastic sediments
should be sought in the northeast. The upper part of the Egerci Formation consists of
lacustrine limestones that are plateau-forming. The rodent locality lgdeli, which is
situated just below the lacustrine limestones, suggests that the base of the upper part
of the Egerci Formation is of Early Pliocene age.
The difference between the sediments south and north of the Elmah Dag thrust
cannot yet be explained satisfactorily because the age of the upper part of the
Dokmeta~ member is not known. Since all of the formation is gently folded (presum-
ably the folding occurred during the tectonic phase responsible for all the SW-NE
striking structures in the area)!.. the top part of the Ceviz<fik Formation cannot.. b~
younger than the base of the Egerci Formation. The mammal faunas from Karaozu
and Kalekoy indicate that sedimentation started again during the Late Vallesian after
the tectonic phase. Since the radiometric dates obtained from the basalts of the
Yenic;=ubuk Formation indicate an age of about fifteen million years before present,
the period of tectonic activity is bracketed. Nevertheless, there are two possible
explanations for the difference between the sedimentary sequences north and south of
the Elmah Dag thrust:
1. The Burtepe and Yeni~ubuk Formations are time equivalent with part of the
Dokmeta;; member and were deposited in a separate sub-basin;
z. The Dokmeta~ member underlies the Burtepe Formation and is thus as a whole
older than MN 4.
THE LOCALITIES
The relative stratigraphic position of the localities from which we have

collected mammals in the Gemerek area could, with the exception of the position of
Horlak 1 relative to Yenikoy, be established in the field (figures 1 and Z). Horlak 1
and Yenikoy both occur above a prominent gypsum deposit. In the section of Yenikoy
this gypsum seems to represent the regressive phase of marine Eo-Oligocene
deposits. In the case of Horlak the gypsum forms the base of the section, so the
underlying strata are not known. There are marked differences in the composition of
the rodent assemblages from Horlak 1 and Yenikoy (figure 3). Comparison with the
European succession suggests that Yenikoy is much older than Horlak 1. It is there-
fore concluded that the underlying gypsum deposits differ in age too. The strati-
graphic position of the two Harami localities (Akljehir area) relative to the sequence
from the Kayseri-Sivas basin can be inferred from their faunal content only. The
assemblages Harami 1 and Harami 3 were collected from one section at levels that are
about 16 m apart. Rodent associations with a similar composition are not known from
64
ORDERS RODENTIA Lagomorpha
r; I
(Super) Dlpodo- I
F'amllloe Muroidea Ollroldoa Saluroldoa CNM-
ilao!JI•- Idea l'.... Ooholonldao
Idea :f
II 0 ~
z] ~ 11
i l l :II
•
~ ll!illllltt!IIJJI!IlltJI l IIll Ij I Ji ~ JI III flIll
ljjdoll ? ~~ ~ ~ !ft E :!
~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~ II IJ
Dondll ~ ~~ ~ ~ ~ ~ ~
~ ~~ ~ ~ ~ ~ ~ j
Karallzll ~ ~ D ~~
~ ~ ~ ~~ ~ ~ ~ ~ ~ ~~
~ ~~ ~ ~
Kaloklly ~ ~~
~~ ~~ ~ ~ ~ ~ ~ "'"'
Gomorok ! ...
~ 0:~ "' z
~ ~ ~~ ~~ 2
Horlak 2 l:i 5 u
~ ~~ ~ ~ c ,.o
Horlak 1b
... I! -
§ -c :E
~~ ~ r·
Horlak 1a ~ :: 1 'ot
~ ~~ ~ ~ 0:
2
... c
Haraml 3 ~~ ~ :a
~ ~ ~~ ~~ ~ t-\: ~ a
li;
Haraml 1 ~ ~ ... 1 -"'"
it
~~
~~
~ ~~ ~ ~~ ~
Yonlklly ~ ~ :ll A
~~ ~~~ 0: II
Fig. 3. Range chart of the genera of Rodentia and Lagomorpha in
the eleven Anatolian assemblages studied. N = Ml + MZ for
Rodentia, N = PZ/ + P3/ + Z x P/3 for Lagomorpha.
Ol
(J1
locanH. . The relative abundance of rodent (super) families and lagomorph.s in percentages
.. lO .. .. .. """'
~ Sciuroldea l::::::::::::::::::::d Dipodoldea
~ Muroldea mJ c.. toroidea [! + !I Geomyoldea
~ Gllroidea D ct.enodactyloidea - I.acomorpha (Ochotonidae)
Fig. 4. Diagram showing the relative abundance of rodent (super) families and
lagomorphs in the eleven assemblages studied. N = M1 + MZ for
Rodentia. N = PZ/ + P3/ + Z x P/3 for Lagomorpha.
anywhere else, but the presence in Harami 1 of Bransatoglis fugax, Glirudinus

euryodon, Vasseuromys aff. rugosus, a species of Mirabella that is more primitive than
the one from Aliveri and Rombach and of two species of Eumyarion that have a more
primitive dental pattern than all other representatives of this genus, strongly suggests
that this association belongs somewhere between Yenikoy and Horlak 1a. The Harami
3 association differs essentially from that of Harami 1 in that it lacks Mirabella and
contains Spanocricetodon. Since Spanocricetodon is considered to be an evolutionary
stage preceding Democricetodon (de Bruijn et al., 1981), it follows that Harami 3 is
older than the "Democricetodon datum plane" (MN 4). One reason for discussing the
age of these faunas in some detail is that preliminary studies of the sporomorph
associations from the Harami section carried out by Dr. Benda (pers. comm.) contra-
dict the stratigraphic position suggested above. His conclusion, namely that the levels
between Harami 1 and Harami 3 contain the Eskihisar sporomorph assemblage,
suggests a correlation to floras from MN 4 or younger. A detailed study of the Harami
section planned for 1989 will attempt to solve this contradiction.
In studyixlg the diagrams of figures 3-5 one should bear in mind that our assem-
blages are very unevenly lpaced Jihrough time. Some assemblages ~~c.~ as the pa~~s
Harami 1 and 3, Horlak 1 and 1 , Horlak Z and Gemerek, and Karaozu and Kalekoy
are very close in age. The picture presented is also very incomplete and therefore
does not allow a detailed comparison of the Neogene fauna successions of Anatolia and
southwestern Europe.
66
LocaiHies The relative abundance of muroid genera in percentages
Oendll
KoroiSzD
Harcml 3
Horoml 1
.. ..
YenlkOy
JO 40 >0 70
[S.S} Sponpcrfc•todon
[I] Crlc.todontlda•
[II} P••udom•rlonn mliJ crlc•fulu•
Cucrlc•#odon Indo!.
IIIIIlD ~ ~ bJ;1 Ot:clfanomy.

m
Pa•udocrlt:.todon Crlc•todon Byzanffnla Paropod•mu•
Cumyarlon 9 IJ~ocrlc•#odon Kowalsklo [}3 Progonomys ~ Apod•mus

O.p•rwfomys ~ O.mocrlc•todon - Blancomy• !IIIIfll Crlcotu• [ill] Al/cromys
ltllra/H/Ia
m Turlcomy• 0 Pllospalax IIJI llesocrlc•fu• [ ! J Prom/momys
Fig. 5. Diagram showing the relative abundance of muroid genera in the eleven
assemblages studied. N = Ml + MZ.
DEFINITION OF INFORMAL ASSEMBLAGE ZONES
It is difficult to evaluate a faunal succession from a poorly known bio-province.

Unless a number of rich associations from each level are available, it is almost
impossible to distinguish between the effects that the age (biostratigraphy), the
biotope (paleoecology), the ' geographical position (paleobiogeography) and the origin of
the accumulation (taphonomy) had on the composition of a given association. It is
therefore clear that the first sequence of faunas of smaller mammals covering a
larger part of the Anatolian Neogene can at best provide the basis for a preliminary
bio-zonation.
Zcme A. The association of Eucricetodon cf. praecursor, Pseudocricetodon cf.

montalbanensis a peculiar derived ctenodactyloid (see figure 6) and a ctenodactylid
that is reminiscent of Asiatic representatives such as Tataromys is considered to be
characteristic.
Zone B. The diversity of eumyarionine cricetids seems to characterize zone B.

We suggest dividing this zone into two sub-zones because the presence of Spanocrice-
todon in Harami 3 seems to record the first arrival of "Miocene" cricetodontids from a
presumably central or southeastern Asiatic center of dispersal. This migration seems
to have had a large impact on the composition of the Muroids (figure 5).
ZODe C. The association of Democricetodon, Megacricetodon, Cricetodon and/or

Turkomys characterizing zone C makes this unit a crude unsophisticated one. The
67
representation of this zone in our collections is poor, but allows the recognition of a
lower part, C 1 , characterized 2 by Cricetodon cf. aliveriensis and Albertona cf.
balkanica and an upper part, C , containing less primitive Cricetodontini. The
presence of a species of Sayimys similar to Sayimys minor in the lower part of zone C
is of ·special interest because similar teeth are known ·from the Lower Manchar
Formation (Indian subcontinent) and from the Xiacaowan Formation (China).
Zane D. The association of Byzantinia as the only cricetodontine and the murids
Progonomys and/or Parapodemus characterizes this zone.
Zane E. The association of such "modern" cricetids as Cricetus, Mesocricetus

and Cricetulus with diverse murids and the arvicolid Promimomys characterizes zone
E.
The Correlation of the Rodent Assemblages
from Anatolia with the MN Zonation
The succession of Neogene rodent faunas in Anatolia evidently has a style of its
own and differs, certainly at the species level, appreciably from the western European
succession. However, when we look at the genus level, and more specifically pay
attention to genera that appear in Europe as immigrants, similarities in the dynamics
of the faunal development of the two areas become apparent. The overall impression
is that the very pronounced endemism of the Oligocene and Early Miocene faunas from
Europe is abolished step-wise by successive migration-waves of Asiatic rodents. As a
result, biostratigraphic correlations between Anatolia and western Europe are
expected to be more reliable in the upper part of the Neogene than in the lower part
of the Neogene.
The association from Yenikoy (Unay and de Bruijn, 1987) is of special interest
because it is the first record of Oligocene rodents from Anatolia. The small collection
discussed in 1987 has been somewhat enlarged during the last summer and consists now
of 42 isolated cheek teeth of Eucricetodon cf. praecursor (i.e., the lower teeth from
Yenikoy. are very similar to the type of praecursor from Quercy), three cheek teeth (1
M/Z and 2 damaged M/3) that seem to represent a larger species of Eucricetodon, the
lingual half of a very large muroid M/Z, one M1/ and one.M/2 of Pseudocricetodon cf.
montalbanensis, one lower dentition of Parasminthus (Unay et al. op. cit.), some
damaged rather bunodont ctenodactylid teeth (cf. Tartaromys?) that are listed as
Ctenodactyloidea indet. in figure 3 and three more or less complete teeth and
numerous fragments of a .J.ophodont semi-hypsodont rodent, that was erroneously
assigned to the Gliridae by Unay et al. (1987). These last specimens (figure 6), identi-
fied as M/1, D4/ and M1-2/ will be discussed in some detail because it is not at all
evident to which family these characteristic teeth should be assigned.
The homogeneity of our sample cannot be established, but even if the M1-2/
would belong to another, smaller, species than the lower teeth, there seems no doubt
that they represent the same family.
The combination of four-crested lower molars with five-crested upper molars,

the sub-equal protocones(ids) and hypocones(ids) in all cheek teeth, the absence of the
longitudinal ridge in upper as well as lower molars and the elongate molariform lower
deciduous fourth premolar are considered to be the major characteristics of our
animal. These characteristics occur in a number of (presumably unrelated) rodent
(sub)families, i.e., the South American Echimyidae, the western European
Theridomyidae, the central Asiatic Ctenodactylidae, the probably ctenodactyloid-
derived eastern Asiatic Yuomyidae, the southeast Asiatic Baluchimyinae and the
African Phiomyidae. Semi-hypsodont four- or five-crested dental patterns developed
worldwide in many rodent families during the Middle Oligocene, a phenomenon that
may well be correlated with the increase of the importance of grasses during the same
period. Among these the Echimyidae need not be considered because of paleogeo-
graphical reasons. Among the Theridomyidae the genera Archaeomys, Issiodoromys
and Pseudoltinomys contain species that, at least superficially, show a dental pattern
68
a
b
b' b"
c
Fig. 6. Cheek teeth cf. Lophibaluchia from Yenikoy. a - D/4 sin. (± z.s x
± 1.8 mm); b- M/1 sin. (Z.37 x 1.90 mm), b'- same specimen anterior view,
b" - same specimen posterior view; c- M1-Z/ dext. (1.87 x 1.90 mm). All
specimens approximately x 25.
similar to that of the specimens from Yenikoy, At this point the interpretation of
homologies of various parts of the cheek teeth enters the discussion. We tend to
consider the often-long central ridge of five-crested theridomyid lower molars as a
true mesolophid. Since there seems no doubt that the four-crested molar-type in this
family developed through loss of the anterolophid, the second ridge of such molars is
the mesolophid. In our specimens the second crest seems to be the enlarged posterior
arm of the protoconid, a situation that is common in ctenodactyloids and the (probably
ctenodactyloid-derived) Yuomyidae, Baluchimyinae and Phiomyidae. It seems, there-
fore, that our specimens have nothing to do with the Theridomyidae. Other arguments
for this conclusion are the straight anterior occlusal outline of the D/4, the absence of
any indication of extra crests in the anterior portion of this tooth, the strong postero-
lophid of the M/1 and the presence of four roots under M/1. None of these features
are known to occur in the Theridomyidae.
This leaves us four (sub)families (Yuomyidae, Baluchimyinae, Phiomyidae and

Ctenodactylidae) as possible candidates to house our enigmatic rodent from Anatolia.
This means, in our opinion, that our animal must have had a ctenodactyloid ancestry
regardless which of these groups (if any) it belongs to. Structurally, the dental pattern
of the specimens from Yenikoy is most similar to Lophibaluchia. Moreover, the type
species of this genus also shows two roots below the anterior portion of the lower
69
molars. Although we think that all the features that suggest assignment of our teeth
to the Baluchimyinae might have evolved in parallel in any other highly specialized
ctenodactyloid, our working hypothesis is that our animal is more closely related to
Lophibaluchia than to any other rodent. A case against this hypothesis is that the
time relationship of the Bugti and Yenikoy faunas is not right. The Bugti beds are
usually considered to be of Late Oligocene or. Early Miocene age while the presence .9f
Eucricetodon cf. praecursor and Pseudocricetodon cf. Montalllanensis in Yenikoy
suggests a Middle Oligocene age. Faunal exchange between the Indian subcontinent
and western Asia prior to the Miocene seems unlikely because of the very pronounced
endemism shown by the rodent fauna from the Bugti beds.
The assemblage from Harami 1 (zone B 1) contains two species of Eumyarion with
primitive dental morphology (strong posterior arm of hypoconid in M/1 and M/Z, strong
anterior arm protocone in M1/), Deperetomys intermedius (de Bruijn et al.)*, a species
of Mirabella that shows strong Eumyarion affinities, Bransatoglis fugax (Hugueney),
Glirudinus euryodon (van der Meulen et al.), Vasseuromys sp., Miopetaurista cf. dehmi
(de Bruijn et al.) and Palaeosciurus costatus (Freudenberg). This association is not
known from elsewhere and can therefore not be correlated directly. The Eumyarion
and the Mirabella strongly suggest that it is older than MN 4 and the absence of
Eucricetodon and Pseudocricetodon indicate that it is post-Oligocene. The Harami 1
fauna is tentatively correlated with MN 1 or z.
The assemblage from Harami 3 (zone B2) differs from that of the older Harami 1
by the absence of Mirabella and the presence of a species of Spanocricetodon that is
quite similar to ~· lii. If our working hypothesis that Spanocricetodon is the ancestor
of Democricetodon is correct (de Bruijn et al., 1981), this assemblage is older than
MN 4. The dental morphology of the two species of Eumyarion present in this associa-
tion supports that conclusion. The Harami 3 fauna is tentatively correlated with
MN 3.
The set of assemblages of Horlak 1a through Gemerek temporarily assembled in

zone C shows (trending upward) a decrease in the number of specimens of Cricetodon
and an accompanying increase in Megacricetodon. The assemblage of Muroidea in
zone C is roughly similar to that in the Aragonian (MN 4 - MN 8) of Spain. The
presence of Cric;.etodon cf. aliveriensis and of the ochotonid Albertona cf. balkanica in
Horlak 1a and 1° suggests that the lower part of zone C correlates with MN 4. The
evolved and larger Cricetodonini present in Horlak Z and Gemerek are clearly younger,
but do not allow a precise correlation with the MN scale.
The assemblages Kalekoy, KaraozU and Dendil contain Muridae that are well
known from the Middle and Late Vallesian of Spain and that have been used to define
formal zones (van de Weerd, 1976). Kalekoy, on lithostratigraphic grounds the oldest
of the three assemblages of our zone D, contains Parapodemus lugdunensis and
Progonomys hispanicus. In-Spain.!:· hispanicus precedes Parapodemus lugdunensis and
the stratigraphic ranges of these species do not overlap. Although our zone D seems
to correlate with MN 10 and 11, precision is again impossible.
The assemblage of Igdeli, though very characteristic, does not contain many
species that are well known in the western Mediterranean. Promimomys insuliferus,
however, conclusively suggests the correlation with MN 14. Among the Muridae there
is a small species of Occitanomys that is known to occur in many MN 13 and MN 14
assemblages, but it has not been formally named [figured in de Bruijn et al. (1970)
under the name Castillomys crusafonti].
*The species intermedius was erroneously assigned to Mirabella by de Bruijn et al.

(1987). New collections show conclusively that it is closer to Deperetomys .than to
Mirabella.
70
Comparison of the Neogene Rodents of
Anatolia with Occurrence Elsewhere in Asia
The Indian subcontinent. Two of the rodents found in Anatolia seem to be

similar to species that were originally described from the Murree Formation in north-
ern Pakistan (de Bruijn et al., 1981). The first is the Spanocricetodon from Harami 3,
a species that is about the same size as ~· lii from the Murree. Its dental morphology
is also quite similar though relative frequencies of some characteristics are differ-
ent. The metalophulid of the M/1 for instance is directed forward in nine and back-
ward in one specimen from the Murree (Z species). In the Harami 3 material two M/1
have the metalophulid directed forward, in one it is backward, in five it is double and
in one it is absent. The direction of the metaloph of the M 1/ from these collections
shows similar resemblances. In the combined material of S. lii and S. khani the
metaloph is transverse in seven and directed posteriorly in one specimen~ In the M1/
from Harami 3 the metaloph is transverse in six, directed forward in twelve and
absent in two specimens.
The second species that seems to occur in Turkey and Pakistan is Sayimys
minor. This species, originally based on scanty material from the Murree, is now also
known from the lower part of the Lower Man~har Formation (Sind). The few Sayimys
specimens collected from Horlak 1a and 1 are morphologically identical to the
Pakistani specimens but somewhat larger than the type material.
Although we are quite aware that diachronism in the occurrence of similar

morphology is likely to occur, it seems of interest to pursue the study of groups that
have a very extensive geographical range. In this context it would be worth studying
the representatives of Pseudomeriones in detail. Our Pseudomeriones specimens from
Karai:Szii are very similar indeed to the specimens from Molayan (Afghanistan)
described by ~en (1983), whereas the material from Dendi! resembles E_. abbreviatus
from King-Yan-fou (China).
CONCLUSIONS
The succession of Neogene rodent faunas from Anatolia shows a number of

differences as compared to the sequence from Europe.
1. Continuous dominance of the Muroidea in all rodent faunas from the Middle
Oligocene onward. The so-called cricetid vacuum (MN 3) does not show.
z. The first arriving "Miocene cricetid" in our sequence is Spanocricetodon and not
Democricetodon.
3. Eomyidae do not seem to have reached Anatolia before the Vallesian, although
they are present in the Middle Oligocene of Thrace.
4. Gliridae are rare, but diverse in pre-Vallesian faunas. From the Vallesian onward
they are better represented in terms of specimens, but the number of genera is
reduced to one, Myomimus.
5. Sciuridae are either rare or absent in the lower part of our sequence, but diverse.
From the Vallesian onward they are somewhat more common, but represented by
just one genus, Tamias.
6. Ctenodactylidae occur as rare elements in many of the pre-Vallesian assemblages

from Anatolia and seem to have been diverse. This family never reached the
European mainland, but is known from the Oligocene of the Balearic island
Mallorca, the Early Miocene of Sardinia and the Pleistocene of Sicily.
71
The Neogene record of the Anatolian Muroidea permits the recognition of five
phases of relatively rapid change in the composition of the assemblages. These phases
are:
1. The appearance and subsequent diversification of the Eumyarioninae. This sub-
family seems to have replaced Eucricetodon and Pseudocricetodon.
z. The appearance of Spanocricetodon, which is supposedly ancestral to Democrice-

todon.
3. The appearance of Cricetodon and Megacricetodon.
4. The appearance of Progonomys, Parapodemus and Pseudomeriones. In our

sequence from the Kayseri-Sivas basin the genera Kowalskia and Pliospalax also
occur for the first time at this level, but they are known from older faunas from
elsewhere in Turkey (iJnay, 1978; Cnay and de Bruijn, 1984).
5. The appearance of Cricetus, Mesocricetus, Cricetulus and Promimomys.
These five events seem to document immigrations of muroid genera into the
area, because in each case potential ancestors are absent from older deposits. The
direct biostratigraphic correlation of the periods of increased faunal turnover recog-
nized in Anatolia with the MN zonation is not always possible because some of the
immigrants never seem to have reached western Europe (Spanocricetodon, Pseudo-
meriones, Mesocricetus) whereas others seem to have appeared there much later
(Eumyarion, Deperetomys, Cricetus, Cricetulus).
As a working hypothesis we propose to correlate the arrival of the

Eumyarioninae with MN 1 or MN z, the arrival of Spanocricetodon with MN 3, the
arrival of Cricetodon and Megacricetodon with MN 4, the arrival of Progonomys,
Parapodemus and Pseudomeriones with MN 9 or MN 10 and the arrival of Cricetus,
Mesocricetus, Cricetulus and Promimomys with MN 14.
If our correlation of the Harami faunas with the early Miocene is correct, the
reconstruction of the geological history of Anatolia will have to be revised because
block-faulting started much earlier than was previously assumed.
REFERENCES
Bruijn, H. de, Dawson, M.R., and Mein, P., 1970. Upper Pliocene Rodentia,
Lagomorpha and Insectovora (Mammalia) from the Isle of Rhodes (Greece).
Proc. Kon. Ned. Akad. Wetensch. (B), v. 73 (5), p. 535-584.
Bruijn, H. de, Hussain, S.T., and Leinders, J.J.M., 1981. Fossil rodents from the
Murree Formation near Banda daud Shah, Pakistan. Proc. Kon. Ned. Akad.
Wetensch. (B), v. 84 (1), p. 71-99.
Bruijn, H. de, Unay, E., Sara~, G., and Klein Hofmeijer, G., 1987. An unusual new
Eucricetodontine from the Lower Miocene of the eastern Mediterranean. Proc.
Kon. Ned. Akad. Wetensch. (B), v. 90 (Z), p. 119-13Z.
~en, ~., 1983. Rongeurs et lagomorphes du gisement Plio.c'Eme de Pul-e Charkhi, bassin
de Kabul, Afghanistan. Bull. Mus. natn. Hist. nat. Paris (C), ser. 5, v. 1, p.
33-74.
Onay, E., 1978. Pliospalax primitivus n. sp. and Anomalomys Gaudryi Gaillard from
the Anchitherium fauna of Sarifay (Turkey). Bull. Geol. Soc. Turkey, (C), v. Z1,
.. p. 1Z1-1Z8.
sides of the Dardanelles. Newsl. Strat., v. 13 (3), p. 119-13Z.
Unay, E. and Bruijn, H. de, 1987. Middle Oligocene to Early Miocene rodent assem-
blages from Turkey, a preliminary report. Miinchner Geowiss. Abh. (A), v. 10, p.
Z03-Z10.
Weerd, A. van de, 1976. Rodent faunas from the Mio-Pliocene continental sediments
of the Teruel Alfambra region, Spain. Utrecht Micropal. Bull. Special Publ. Z, p.
1-Z17.
72
UPDATING OF MN ZONES
P.Mein
Departement des Sciences de la Terre

27-43 Bd. du 11 Novembre
696ZZ Villeurbanne Cedex, France
The recent numerous advances in our knowledge about fossil mammals have
required a review and revision of the MN zonation (Mein, 1975, 1976).
In some areas (especially in Spain) the succession of biological events among

Cenozoic mammals can be identified in a very sharp manner, especially with the
evolutionary grade method of some rodent lineages.
Unfortunately, small mammal species generally have a limited geographical

distribution that does not allow long distance correlations. So in this new chart,
biological events are mentioned only at the generic level with special references to
faunal records of west and central Europe.
The revised chart of MN zones, common taxa, generic first appearances in

Europe, and generic last occurrences in Europe are shown in table 1. Table 1 follows
the format of previous charts of MN zones (e.g., 1975 and 1979) with numerous addi-
tions and slight revision. MN zones are grouped in table 1 according to mammal ages
of Fahlbusch (1976). Another notable change is that the designated reference fauna
for each subdivision is printed in capitals. The most important biological event is the
first appearance of ·a new genus, but we also note the last occurrences.' For each
mammalian age, the list of the main genera are also given.
To allow greater ease of correlation we have tried to simplify the subdivision

and have retained formal subdivision (a and b) only for MN z.
For the same reason, we have not split the last Neogene unit MN 17 (corre-
sponding to MN 17 + 18 of Guerin). With the new recognition of the Neogene-
Pleistocene boundary at the top of the Olduvai event, correlated with the beginning of
the Eburonian cooling, the majority of paleontologists now place within the Neogene
some faunas, like Seneze, which were formerly considered Pleistocene.
In this revision, we have tried to keep the change of localities position to a

minimum and have added several new and well studied faunas.
For the Agenian age, there are no major modifications, only the addition of new
Spanish localities.
For the Orleanian age, the French locality Artenay is placed again in MN 4; that
determination is based on the new record of Democricetodon (Bulot, oral communica-
tion). So, as previously argued by Ginsburg, the Proboscidean datum is placed at the
beginning of MN 4 (for western and central Europe). During this interval the biolog-
ical events are so numerous that it was impossible to note all the changes.

-..I
....
Table 1. European Mammal Neogene (MN) Zones.
AGENIAN
Common Taxa rlnt Appearance Last Occurrence

loiN W. Europe C. Europe E. Europe S.E. Europe W. Asia N. Africa
W+C Europe Other W+C Europe Other W+C Europe Other
lorane a Amphipora-
Bouzlguoo thorium Palaeo:scaptor Lagopsls Tltanomys
Navarrete ~~~r.~h Plozoduo Agyopolaguo Prologuo Plozoduo
Heaaler Amphllaguo
LAUGNAC Tltanomyo Brolllana Horpostldos
Ploologalo Proallurua
2b Schaff- Eucrlcotodon Parataplruo
Solloo-our-Chor hauaen Paeudocrtce- T~!1:l:.rr•- Xonohyuo Pleuroceroa
todon Dromothorlum
lolelloolodon ~~~~::;:. Aurollachooruo lllrobolla
Ballzac Stonooflbor Protalactaga Depere-
tom yo
Haraml 1 Cynoloe Eumyarlon
--- ------ ------- Warculnomya -----------
Cocumant La Chaux
------- Amphlcyon RIHonorla Rhodanomyo
Valquomado Haplocyon Armantomya
Marcotn Cophalogolo Pooudodryomyo
Yoongrlnla Horpootldoo Proolluruo
Lambert Ploolctlo Potamothorlum
2a Chavrochoo Amphlctlo
Colina Polaoogalo Phylotlllon Protaplruo
IIONTAIGU Orlomoryx Palaeochoerua
Alllao
Gano Agyopo Dlacorathorlum
lololooac Welaenau
----- --------- ------- -------- Proto cera- Paracera- Dlmylechlnuo
----------- ------------
thorium thorium
RoHonburg Aprotodon Tltanomyo
Saulcot 3,5,8
Apoomyo
Wolooon- Amphlcyon
1 loloncalvlllo burg 6 Hyotherlum Cyneloo
Palaoogalo
rornant 11 Calnothorlum lollomoryx
Lophlomoryx Proatlurua Eotaplruo
PAULHAIC Tomordlngon Dromothorlum Prodremo-
thorium
Wlochberg Amphltraguluo Hyotherlum Bedenomeryx
Elomeryx
Pyrlmont-
Challomyo
Boudry 2
ORLEAN IAN
Common Taxa First Appearance Last Occurrence

loiN W. Europe C. Europe E. Europe S.E. Europe W. Asia H. Africa
Qta Pombolro Leo ben Chloo Prolagus Plloplthecus Amphlporalherlum

Pontlgno Puttenhauoen Slbnlca Yeroham Lagopolo Llgerlmyo
Chela• Savlgno Maaaendorf Rotom Eomuacordlnua Turkomyo Palaeoaclurua
t.lunebrega Langen- Spermophlllnus Keramldomys Anchllherlomyo
moosen AI Sarrar Anchlthorlomys
AI Dabllyeh Crlcetodon
Toffalba Thonay Oggenhof Komollnl P11udaeluruo Chollcotherlum
5 PONT LEVOY Koylbulax Hoskoy Horpootldos Chllotherlum ~::~::Y~!~Ium
Qta Pod relzas franzenbad Galndatherlum Hlspanotherlum
Vermeo 1 t.lurefte ~~";~,:~:~a
Luzovlk Ursovua Bunollslrlodon
Boaugency Elblowald
Tarazona t.lala t.llllva Conohyuo Aurellachoeruo
lavers Sanlthorlum
Vloux Collongos Strakonlco Bania Luka Gemorok Dlcroceroa Sanlthorlum
Heteroprox
t.lonlroal ------ ----- ------- ----------
Gobel Zollon Anchlthorlum Ptychoprolaguo
-·---------- lofolloolodon
Alberlona
---------
Forothart
Qla Farlnholra Rombach Oemocrlcetodon Pooudolherldomyo
Balgnoaux Erketohofon ~~T~~~~~:~um Megacrlcetodon Antomuo Cynoloo
t.lorallneo Aorotroln Orechov Anomalomya
Rou1cherod ~:?:!:~'!~x Neocometee Shamallna Protacerothertum
4 Co reo lea Folnln Albanohyuo Hyoenallouroa Araboa- Olacerotherlum
LA ROt.IIEU Blshlubo Horlak 1 mlnthus
AIIYIM Prodolnothorlum ~:~T!~C:~:therlum Crlcetodon
Proaantorhlnua Brachyoduo
Gompholhorlum Larlotothorlum Procervulua
Bunol PROBOSCIDEA Bunollo-
Valtorea Zygolophodon Bunollotrlodon trio don Amphltraguluo
Arlenay Dol nice Jabal t.lldra Eotrogus Canthu-
Qta Narlgao Potorobuch Archaoobolodon Mtcromeryx meryx
Kln!vtoh• -- .LWo_t.ltcu._ Oorcatherlum Dorea-
Moratllla NOUYIIIO Tuchorlco --- ··················· Stephanocemoa thorium
Wadi Llgerlmys
-------- Amphllaguo
----------
Chlllours Schnaltholm sr::::rlce
t.loll Calopa WINTERSHOF- t.loghara
WEST Eucrlcotodon
Serre de Verge• Ureavua Paeudocrlcetodon
Agroda Welaaen- Pseudarctoa
3 burg 5 t.llrabello ~~:~~r.:~~~~hllua
Cootablanca Haraml :S Eumyarlon Anchlthorlum
Deperotomyo Amphlmoschuo
Valhondo Chomatov Xonohyua Paralaplruo
Estropouy Echzoll Brachyoduo Xonohyuo
Bluing on Lagomeryx Andegameryx
Hona du J~:r."n~Y Acteocemaa Orlomeryx !
Unlv. Colo. Llsboa ~~y;~~urg Stephanocemaa
Leo Bolllaux Procervulua
-.1 (continued)
C1l
.....
en
Table 1 (continued)
ASTARACIAN
Common Taxa First Appearance Last Ocourronce

loiN W. Europo C. Europe E. Europe S.E. Europe W. Aala N. Africa
W+C Europe Othor W+C [uropo Othor W+C Europo Othor
Plloplthocuo Dryopllhocua Pleslosorex
Yen I Crlcetulodon Crlcolodon t.le;apodotoa
San Qulrzo Eoklhlaa Galorlx Ooomanolla Doperetamyo
La Grlvo L3 Lanthanothorlurr Cruoafonllna Zramyo
Kloln- Palanlak 6 Plotlodlmylua Konyalagomyt Wuscardlnus Neocomete1
B Excobooa oloonbaoh
St. Gaudono Vormeo 2 Pneworno Boyrok- Prolaguo Alloptox Hlopanomyo
Glv;enhouoon II lepo 3 Lagopala
Solera lolelaaaylmya
~~~ 1 Etephan Crlcelodon Ko:1;nochoerua Palaeomeryx
La Clotornlga Sofca Afrlcanomya Pro ragocerue
lufJJ.U'______ lolegacrlcetodon
Domocrlcotodon lologaflodotoa Tolralophodon Plalybalodon
1~;ro-au lochyrlctla Proia lactaga Dolnolhorlum Prodolnothorlum
----- ---------- ---------
Palencia Lodo ---------- ----------
La Grlvo lol ~:~t·!~:r~~~=
Sansano1mllu1
Corro dol Otoro STEIHHEIIol Sarlcay roraythla Dakkamya Hyaonallouroo
7 Oppolo Anchllhorlum Homlcyon
La Grena11ere Plackla Benl lolollal Protlcllthorlum Olnocyon
~~:~~f::~r:~~" Paradlcerow ~TI~~~~r~= Plltoayon Hlapano-
Torll Slmorro Plealaceratherlum therlum
Koroth.l Larlelothorlum Mesaceratherlum
Ohnlngen lollkulov Ph;ylotlllon Euprox
---------- ------------ --------- ----------- Challcothorlum
Paracuolloo 3,5 -·---------- -----------
Wanchonea Goorgono- Slvapllhocuo Doamanodo• Ptychoprolagua
mund Candlr Crouzella Schizo-
Hyothorlum
Arroyo dol Val Hasno1 Conohyua turolagua "''~~i~~ Gllrudlnuo Turkomyo
Sandolz- Albonohyuo Taxodon
hausen Dorcathorlum lollohyaona Goblcyon Pooudocyon
Szontondrol Logomoryx Parcrocuto Tungurlctlo
6 Uot lollcromeryx Percrooulo Colnotherlum
Kloln AI Jadldoh Olcrocerue Taucanamo
Hadorodorf Pr•breza Palaeomeryx Kubano- Dromacara- Bollalovlna Coucaso-
Stolnborg Eotrav~• choerue thorium thorium
Gclldborg Kubanotragu• Aceratherlum
SAN SAN Schononborg Prodolnothorlum lnlanathorlum Llotrlodon_ Llotrlodon Archcooboladon
Gerlach Blolo-
Noudorf metschav Paoolar Zygolophodon Plalybolodon Capralragoldeo _ Capra-
Sandbo'l, tragoldoo
Houdarf p. Gomphothorlu m Choero- Hypoodonluo
lophodon
VALLES IAN
Common Taxa Fltot Appearance Laot Occurronco

liN W. Europe C. Europe E. Europe S.E. Europe W. Aola N. Africa
Dendll
Cucalon Kohfldloch Kalnary Cruoafontlna Parapodomuo Alllopuo Eurolaguo
Amborlou 1 Lofkon Amama 1 Eplmorlonoo Lopho- llogacrlcotodon Afrlcanomyo
Suchoma1ty Galorlx Eozapu1 crlcotu1 llyogll•
Torraooa Groooulova
Pralaguo Proochotona
La Tarumba Agha Jarl Ouod Zro Sarmatamyo
Sob lay Ravin doo Hlopanomyo Byzantlnla =~~~~1::r:ta Uroavuo
10 Borlolav Zouavoo(RZ1) Crlcotulodon Zramyo Alloopalax Pooudo-
Bladrak Kobaahta lluocardlnuo llyocrlcotadon merion•• Sanoanoomlluo
llontrodan Gotzendorf Karaozu Spormophlllnu• Poeudaeluruo
IIASIA DEL BARBO Eldar Koudlct Protalactaga
ot Tluo Albanonola
Adcrocuta
loflapotaurlota Ouzoceroe Schlzochooruo
Bohlin Ia
Kalakoy Tragonthorlum Adcrocuta Honano- lollatragaceruo
Lyon X Rouooo Corak thorium
Vooondorf Balclk Yorlor Palaoomyo Samotraguo Euprox
VIlla do Caval• Karaln ht•••m-
Ravin do Ia A Kin lofachalroduo brlacoruo
Plulc Bayrak- Thalaoalcllo Tragoportax Prootrop-
_!!p_o.J!_ Lycyaona tiS:.!1!9.!__
-- Kaotollloo-- ~----- Micro ocr ce ue ~1no1orex
----------
lndarctoo ~~rca:.~; lofotacordylodon
Jobol Somono Progonomyo Progonomyo
CAN UOBATERES Galsolborg Sobaotopal lofahmut- Pllavlvorrapo Plloplthocus
Aumel1ter lalovena kay Bou Uroavuo Pllopeteo Cryaplthecu1
Can Ponslc Grooolappon Hovorany Hanlfla5 Zramyo
St. Joan do B. Vamltoa Challcothorlum Sonouoolmyo
Podreyguoros 2C Charmolllo Elmo Bou Taplruo Buzhoromyo Domocrlcotodon Dakkamyo
Akcakoy Hanlfla 1 Acorathorlum Chllathorlum Eomuacardlnu•
Schizo- Eumyarlon
Lapuohna Korynochoeruo galerbc Agnotherlum
9 Rudabanya Dorcathorlum IProttctllhorlum
Lao Valloo do Atavaoka loflcromoryx Samo- Amphlcyon Mlomach-
Fuontlduona Hammer- thorium Trocharlon alroduo
achmlodo Broil a Yaoolorom ~~ra'::traguo loflcrootonyx Llotrlodon Dlcorypho-
Carrllanga lollotragocoruo Lagomoryx choeru•
Douo Ia Fantalno llarktl Kalfa Totralophadan Plio hyrax Dtcrocero•
Blod Douarah Chooro- lndarctoo
AzambuJolra 1 Howonogg Buzhor lophadon Mochalroduo lofacholroduo :r:!~o\;~".ru•
Nombrovllla ~r:~:::r~~::.n Dlcorothlnuo Dlcoroo
JuJurloux Comanutl 2 Oryctoropuo Hlpparlon Hlpparlon Anchlthorlum
Can llata 1 Eppoloholm Gomphothorlum
-.J (continued)
-.J
-..!
(X)
Table 1 {continued)
TUROLIAN
Common Taxa rtrst Appearance Lao! Occurrence

MH W. Europe C. Europe E. Europe S.E. Europe W. Asia H. Africa
Arenaa del Roy
Brlolgholla Mooopllhocuo Wacaca
Salobrona Sahabl lollcrolodon Adcrocuta
Angustovka Wadi Hatrun ~:.::~:::::.~~~ Analolomya Thala01lctls
Ubrllla Khondok Galorlx Hyolrlx Mlcroto- Wyocrlootodan
13 ol Oualda Schlzogalerlx Apodemua acopt11 Acoralhorlum
Crevlllenlo 6 Taakln- Diboll a Paraolhomya lachymomya
Caolno Dlllko paoa Doamanolla Crlootua
Vonla dol lotoro Ballavar Mamay Monaolorl Anouroaorex Cervavltua Mlcroatonyx
Ano Wotochl Amblycoptua . .~:~.r::r:. Pllocorvuo Palaooryx
EL ARQUILLO Bacclnollo V F'ranzovka Blarlnolla Hollodothor1um Palaeoreae
:·C~aamapor1hotoa
La Por1ora Hatvon Dlnofolla Rodunca Dolnolhorlum Choorol-
LIIIIOU Polgardl Agrlolherlum Paraoamolu Tetralophodon phodon
lotollna do Sequra Samoa 5 Wenacer Prolagua Paracamelua
Tudorovo Plkorml - -
Loa Al)ozareo Amblycoptuo
------- ----- Hlopanomyo Dlnoaorex
Choma tori Ruaclnomya Ruactnomya ~r:~'~y. Ploalodlmyluo Zromyo
LOS MANSUETOS Tardoabanya Kowalakla Stephanomyo Byzantlnla
Salonlkl lhroudla Hlapanomyo
Cucuron Slade Paropodemuo Crlcetu-
12 Wr. Leberon Klnlk Occltanomyl Protatoro Ioden
Bucclnollo VI "Valorymyo" Pooudo- Percrocuta
Con cud Cobrucl Amama 2 Eozapua merton••
(Cerro do Ia Garita Ellomya Selene-
Clmlslla Atlantoxerua Hlopanodorcoa Mlcromoryx
pf.~~~~UI
Casa del Acero Taraklla ~f:;:,~g=~~~· Gazella
Tlhana ----- --
------------- Ifoli"iir.lF --------- ~:r..~~~=~ropo Anacue
Lobrlou Novo Samoa 1,4 gozl
Amberlou 3 Elloavotouk Adorocuta Paramo- Lanthanothorlum
llollon Elchkogol Garkln cholrodu• ~~~r.:':amy• Slvapllhocua
La Cantero Hlpparlon "Valorymyo" Anomalomyo
St. Bouzlle Sumeg Corak Challcolhorlum Dlpoldto Alloopalax
Proapola~
Dorn- Yel•r Dlcerorhlnus Dlceros
Durkhelm Kucuc Sldl Solem
11 PI ora Cekrace lllcrostonyx Toplrlscuo Crlothorlum Ooconnalhorlum Samothorlum
Bacclnollo VC Poraholy I.Moravha Gazella Cervovltua Ploaladdax Plosladdox
CREVILLENTE 3 Raopopeny Prochoma Tragoportax Orycleropue Psoudotraguo Palaeorea1
Dlonoy Stogototra- Protoryx Hlsldorcaa Hlsldorcaa
Coakvar Grobenlkl Vathylakkos Kayacllbl bolodon Holladothorlum
Ravin dos Delnothorlum Choorolo-
Zouavoa(RZ5 Zygolophodon phodan
RUSCINIAN
Common Taxa nrst Appearance Last Occu.,..nco

liN W. Europe C. Europe E. Europe S.E. Europe W. Asia N. Africa
Mooopllhocuo
Layna Wolforoholm Boremondla Galorlx
~=~:~~~OCUI
plthoCUI Oryctolaguo
Macaca Pllopentalaguo Ochoto- Trlochlzolaguo
Calla noldoo
Gorafo 2 Solo Coamota 2 Wozo
Aln Brlmba Blarlnokloo Condylura Kowalokla
Orrlon 1,7 Spllla 2,1 Diboll a Mlmomyo Vlllanyla Occltanomyo
~::rz~~~~~~. Borsodla Paraothomyo
Dolomyo Coorla
~~':.\\~:Iague Crlcotuluo Synaptamyo Dlnofollo
C.'J1:~~a A~ta lvanovoe Oryctolagus Scluruo Lemmue
15 Porplgnan Hypolaguo Paralluruo Aratomyo
(Serra! don Kagul Mooocrlcotuo
Vaoquor) Plolomolo I Ruoalnomp EplmotJonoo
La Juliana Odoooa
(Cata- Stophanomyo Poeudo- Wlcroapalax
combo) Caotlllomp merfone1 Pllaoolo-
La Gloria 2 Borootl, lchkoul Paraothomyo lhrondla vlnla
Maluotonl Occltanomp Eulmuo Poracenulua
Cluporcon Plloopalax Orycloropuo
~h=~:::,o:,~•mus
Prror,otaurlota Dallcho- Paranou.:-- "Protatera" Pr"OiiCiiiiiOna-
or roeorex Spermaphlllnuo Amblycapluo
- - - Cao
------- ---- Argoub
VIllalba Rio Go dolo Kutchurgan
Wont Helene Hyotrlx af:::~ne.:l~:. Promlmomyo
Kardla Eploarlculuo ~~r,,o,!,;~uo Coladonola Lopho-
La Calera Ootramoo 1 Kamonokoo Nycloroutoo Arctomolos Promlmomyo Weao- crlcetue
Canto Baranogalo Trllophomyo crlcetua Micro-
Orrloo 1 Antlpavka Chaomapor- Eolramomy toocoplo•
thole• Pachycrocuta Proopalax
Pochycrocuta
Ursuo Moloo Vormola Ploologulo
Colada• I Torrato Cugonouka Ouod Pllovlvorropo
14 Ootramoo Athmonla Hlpparlon
Vondargu.. 1,11 Plolomalo 1 lgdoll Dlcororhlnuo
La Gloria 4 Frunzovka Taplruo
Bacclnollo
Wantpolllor VI Dorkovo Suo
Caravaca Dinar Crolz:etoceroa Slvathorlum
Aklaky Para boo Suo I
Alcoy
Hautlmaen• Gazella Paracervulue
PODLESICE Warltoa
Collonouvo AIn Zygalophadon Stegotetra-
PoraloJoiJ' Kaoolaklno Cuottura bolodon
Hautorlvo Anaj'ICUI
(continued)
....
(I)
()')
0
Table 1 (continued)
"VI LLAr\YIAN"
Common Taxa First Appearance Last Occurrence

MN W. Europe C. Europe E. Europe S.E. Europe W. Asia N. Africa
Allophalomys
Q Brielle Botfla XIII Hippopotamus
--~----- ------------ 1------- ------- --------- ----------·
t.tontou••• 5 Tegeler Kadzielna Paradollcho- Pllolraguo
Dhobol pllhocuo Llbralcos Gallogoral
Oolramoo 3 Dragoneoll Orouaae Dollcho- lotegalovlo Procamptocerua
Seneze r/clthocuo Anancua
Chlllac acaca
Lo Coupe! Chorlootll
De1mana Lagurodon Stophanomyo
Puobla do Valverde Khapry Bulla Rogla Blarlnoldos Clothrlonomyo
Beremendla Lemmua Parallurua
17
ST. VALLIER Llventoovka Prolaguo Vulpeo Chaomopor!hetoo
Oryctolagus Lynx
Hypolaguo Hlpparlon
Pardlnoo Slallna 2 Eucladocoroo
Vlllaroyo
Roccaneyra Clupercenl ~::;:~r: Heaperldocerua Gazella
Pllomyo
Valdeganga 1.11
------ -----------
Cornateotl Amama 3
----------- -----------·
Caotlllomyo
~r;,o.,:~therlum Vlrotalluruo
Tuluceotl Tourkobounla
Etoualrea Montopoll Hyotrlx
El Rincon
Seyne1 Roblellce Gulyazl Nycteroutoo
Oolramoo7 Canlo Equuo
Pachycrocuta
Woreda Kvabobl Aclnonyx Mammuthu• Zygolophodon
r-------------- -------- Limn I Meganthereon
16 Beremend 5 Caponi Homotherlum Paradollcho-
Dlcerorhlnuo Trllophomyo
Hajnacka Uryv 1 Taplruo Kr.lll:::~~· Bloncomyo
Coorla
Balaruc 2 Arclllo Suo Meganthereon Dlnofollo
Gazellooplra Paracamelu1 Homolherlum
VIa Ieite ARONDELU Pllotraguo Aclnonyx Agrlotherlum
Crolzetoceroa Leptoboo
Eocorlhuela TRIVERSA Pllolraguo
Ana neue Gazellosplra
Table 2.. Principal reference localities and fauna.
MNl
F- 47 Paulhiac
Marsupialia Amphiperatherium frequens
lnsectivora Amphechinus arvernensis; Paratalpa micheli; Clapasorex
bonisi
Chiroptera Nyctinomus stehlini; Myotis sp.
Lagomorpha Piezodus tomerdingensis; Titanomys visenoviensis
Rodentia Eucricetodon collatus; Peridyromys murinus; Bransatoglis
concavidens; Bransatoglis fugax; Glirudinus glirulus;
Heteroxerus paulhiacensis; Heteroxerus lavocati; Rhodanomys
schlosseri; Pseudotheridomys parvulus; Plesiosminthus cf.
myarion
Carnivora Cephalogale ginesticus; Cephalogale gracile ursinus;
Haplocyon elegans; Haplocyon crucians; Haplocyonopsis
crassidens; Amphicyon cf. astrei; Ysengrinia tolosana;
Amphicyonine sp. 1; Amphicyonine sp. 2.; Plesictis solidus;
Plesictis palustris; Plesictis' cultellatus; Plesictis sp. 1;
Plesictis sp. 2.; Proailurus lemanensis
Perissodactyla Paleotapirus cf. d.ouvillei; Brachypotherium lemanense;
Aceratherium paulhiacensis; Diceratherium pleuroceros;
Dicerorhinus cf. minutus
Artiodactyla Hyotherium major; Cainotherium geoffroyi; Dremotherium
feignouxi; Amphitragulus major; Amphitragulus cf. lemanensis
MNZa
F - 03 Montaigu
Marsupialia Amphiperatherium frequens
Chiroptera Hydromops stehlini; Palaeonycteris robustus; Rhinolophus
lemanensis
Insectivora Amphechinus edwardsi; Dimylechinus bernouillii; Dimylus
paradoxus; Oligosorex antiqrms; Soricines 1, 2.
Lagomorpha Titanomys visenoviensis; Piezodus cf. branssatensis;
Amphilagus cf. ulmensis
Rodentia Eucricetodon gerandianus; Rhodanomys schlosseri;
Pseudotheridomys parvulus; Plesiosminthus myarion;
Pseudodryomys aljaphi; Heteroxerus paulhiacensis;
Palaeosciurus feignouxi; Steneofiber eseri; Peridyromys
murinus; Vasseuromys priscus; Vasseuromys rugosus;
Microdyromys sp.
Perissodactyla Paleotapirus douvillei; Diaceratherium pleuroceros;
Protaceratherium tagicum; Mesaceratherium paulhiacensis;
Diceratherium aginense; Moropus sp. or Phylotillon sp.
Carnivora Pseudocyon crassidens; Cynelos rugosidens; Cynelos
lemanensis; Haplocyon elegans; Haplocyon crucians;
Pseudocyonopsis gerandianus; Potamotherium valetoni;
Cephalogale depereti; Cephalogale gracile; Stenogale julieni;
Stenogale brevidens; Proailurus lemanensis; Plesictis
lemanensis; Plesictis? croizeti; Plesictis julieni; Plesictis
stenoplesictoides; Plesictis robustus?; Plesiogale angustifrons;
Palaeogale minuta; Paragale hurzeleri; Amphictis lemanensis;
Amphictis antiquus; Herpestides antiquus
(cont.)
81
Table Z (continued)
Artiodactyla Hyotherium major; Palaeochoerus tyPus; Dremotherium

feignouxi; Amphitragulus elegans; Amphitragulus lemanensis;
Oriomeryx major; Amphitragulus boulangeri; Amphitragulus
gracilis; Cainotherium commune
Pholidota Necromanis sp.
MNZb
F - 4 7 Laugnac
Marsupialia Amphiperatherium freguens
Insectivora Proscapanus primaevus; Carposorex sylviae
Chiroptera Myotis insignis
Lagomorpha Prolagus cf. vasconiensis
Rodentia Eucricetodon aquitanicus; Melissiodon sp.; Peridyromys
occitanus; Peridyromys brailloni; Vasseuromys rugosus;
Glirudinus cf. modestus; Steneofiber eseri; Pseudotheridomys
parvulus
Carnivora Haplocyon elegans; Haplocyonoides mordax; Cynelos
rugosidens; Cynelos lemanensis; Amphicyon cf. astrei;
Ysengrinia sp.; Plesictis aff. solidus; Plesictis laugnacensis;
Palaeogale minuta; Plesiogale angustifrons; Amphictis
aginensis; Herpestides collectus; Proailurus lemanensis
Perissodactyla Palaeotapirus cf. douvillei; Diaceratherium aginense;
Plesiaceratherium platyodon; Diceratherium cf. pleuroceros;
Diceratherium sp.; Protaceratherium minutum
Artiodactyla Xenohyus venitor; Hyotherium major; Cainotherium geoffroyi;
Cainotherium commune; Dremotherium feignouxi;
Amphitragulus gracilis; Oriomeryx cf. major; Andegameryx
andegaviensis
MN3
D- Wintershof-West
Marsupialia Amphiperatherium freguens wintershofense
Insectivora Heterosorex neumayrianus subseguens; "Sorex" pusilliformis;
"Sorex" stehlini; Soricella discrepans; Paenelimnoecus
micromorphus; Cordylodon intercedens; Plesiodimylus
hUrzeleri; Talpa sp.
Chiroptera Rhinolophus aff. lemanensis; Rhinolophus sp.; Megaderma sp.
Lagomorpha Prolagus vasconiensis; Amphilagus ulmensis
Rodentia Ameniscomys selenoides; Plesispermophilus decedens;
Paracitellus eminens; Heteroxerus wintershofensis;
Palaeosciurus fissurae; "Ratufa" obtusidens; Miopetaurista
dehmi; Pseudocricetodon sp. ?; Melissiodon dominans;
Pseudodryomys ibericus; Pseudodryomys gregarius; Glirudinus
modestus; Glirudinus gracilis; Heteromyoxus schlosseri;
Bransatoglis spectabilis; Miodyromys biradiculatus;
Peridyromys murinus?; Ligerimys lophidens; Ligerimys
antiquus; Apeomys turkheimae
Carnivora Cynelos rugosidens schlosseri; Cynelos lemanensis helbingi;
Amphicyon steinheimensis bohemicus; Megamphicyon
giganteus; Pseudocyon sansaniensis aff. intermedius;
82
Pseudarctos bavaricus dehmi; Phoberocyon huerzeleri;
Hemicyon sp.; Hemicyon dehmi; Ursavus elmensis; Plesictus
aff. pygmaeus; Plesictis aff. sicaulensis; Plesictis mayri;
Plesictis vireti; Plesictis humilidens; Amphictis aff. antiquus;
Martes laevidens; Laphyctis vorax; Laphyctis comitans;
Palaeogale minuta; Palaeogale hyaenoides; Plesiogale
postfelina; Broiliana nobilis; Stromeriella franconica;
Miomephitis pilgrimi; Semigenetta elegans; Progenetta
praecurrens; Pseudaelurus transitorius
Perissodactyla Anchitherium aurelianense; Tapirus aff. intermedius;
Rhinocerotidae indet.; Mesaceratherium aff. simorrense;
Protaceratherium minutum; Diaceratherium aurelianense
Artiodactyla Aureliachoerus aurelianensis; Cainotherium bavaricum;
Procervulus dichotomus praelucinus; Amphitragulus cf.
boulangeri; Palaeomeryx aff. kaupij Andegameryx serum;
Lagomeryx parvulus
Pholidota N ecromanis franconica; N ecromanis parva
MN4
F - 33 La Romieu
Insectivora Galerix cf. exilis; Talpa sp.; Proscapanus sp.; Heterosoricinae;
Miosorex grivensis; Soricidae indet.
Chiroptera Hipposideros collongensis; Myotis sp.
Lagomorpha Prolagus aff. oeningensis; Lagopsis penai
Rodentia Megacricetodon cf. collongensis; Megacricetodon bezianensis;
Democricetodon mutilus; Democricetodon aff. romieviensis;
Democricetodon sp. 1; Democricetodon sp. 2.; Eumyarion cf.
weinfurteri; Melissiodon cf. dominans; Ligerimys antiquus;
Ligerimys florancei; Peridyromys aquatilis; Peridyromys
occitanus; Peridyromys gregarius; Miodyromys biradiculatus;
Pseudodryomys ibericus; Glirudinus modestus; Microdyromys
koenigswaldij Microdyromys sp.; Bransatoglis cadeoti;
Miopetaurista sp.; Heteroxerus rubricati; Spermophilinus
bessana; Palaeosciurus cf. fissurae; Steneofiber depereti
carnutense
Fissipeda Megamphicyon giganteus; Pseudocyon sansaniensis;
Semigenetta repelini; Pseudaelurus quadridentatus
Perissodactyla Anchitherium aurelianense; Brachypotherium brachypus;
Plesiaceratherium lumiarense; Lartetotherium sansaniensis;
Prosantorhinus germanicus
Artiodactyla Aureliachoerus aurelianensis; Bunolistriodon lockharti;
Hyotherium soemmeringi; Cainotherium miocaenicum;
Dorcatherium cf. naui; Procervulus dichotomus; Lagomeryx
d. rutimeyeri; Palaeomeryx kaupi
Proboscidea Gomphotherium angustidens; Deinotherium cuvieri
MNS
F- 36 Faluns Pont Levoy- Thenay
Primates Pliopithecus piveteaui
Insectivora Plesiodimylus sp.; Lanthanotherium sp.; Soricidae
Lagomorpha Prolagus oeningensis; Amphilagus ulmensis
(continued)
83
Table Z (continued)
Rodentia Spermophilinus besana; Miopetaurista cf. lappi; Miodyromys
cf. aegercii; Steneofiber depereti; Cricetodon aureus;
Megacricetodon collongensis; Megacricetodon lappi;
Democricetodon mutilus; Eumyarion weinfurteri
Perissodactyla Chalicotherium grande
Artiodactyla Procervulus dichotomus; Dicroceros elegans parviceros
MN6
F- 33 Sansan
Primates Pliopithecus antiquus, Crouzelia auscitanensis
Insectivora Dinosorex sansaniensis; Paenelimnoecus crouzeli; Miosorex
desnoyersianus; Oligosorex prevostianus; Blarinella robustus;
Talpa minuta; Alloscapanus sansaniensis; Proscapanus sp.;
Mygalea antiqua; Lanthanotherium sansaniensis;
Lanthanotherium tobieni; Pseudogalerix exilis; Galerix sudrae;
Amphechinus ginsburgi; Plesiodimylus chantrei
Chiroptera Myotis murincftdes; Myotis elegans; Eptesicus noctuloiiies;
Eptesicus campanensis; ?Miomegaderma gaillardi
Lagomorpha Prolagus oeningensis
Rodentia Miopetaurista sansaniensis; Blackia parvula; Spermophilinus
bredai; Heteroxerus cf. grivensis; "Sciurusn sp.; Cricetodon
sansaniensis; Democricetodon gaillardi; Democricetodon
crassus; Megacricetodon minor; Megacricetodon gersi;
Eumyarion medium; Eumyarion sp.; Eomuscardinus
sansaniensis; Myoglis meini; Paraglis astaracensis;
Microdyromys miocaenicus; Miodyromys aegercii;
Keramidomys octaviae; Steneofiber sansaniensis;
Trogontherium minutum
Carnivora Amphicyon major; Pseudocyon sansaniensis; Pseudarctos
bavaricus; Alopecocyon goeriachensis; Hemicyon sansaniensis;
Plithocyon armagnacensis; Schlossericyon viverroldes; Martes
sansaniensis; Martes muncki; Ischyrictis zibethciides;
Proputorius sansaniensis; Taxodon sansaniensis; Mionictis
dubia; Semigenetta sansaniensis; Viverra medica;
Stenoplesictis aurelianensis; Pseudaelurus quadridentatus;
Pseudaelurus turnauensis; Sansanosmilus palmidens
Perissodactyla Anchitherium aurelianense; Chalicotherium grande;
Aceratherium tetradactylum; Brachypotherium brachypus;
Lartetotherium sansaniensis
Proboscidea Gomphotherium angustidens; Deinotherium laevius
Artiodactyla Listriodon splendens; Taucanamo sansaniense; Dorcatherium
crassum; Dicroceros elegans; Heteroprox larteti; Micromeryx
flourensianus; ?Strogulognathus sansaniensis; Palaeomeryx
magnus; Eotragus sansaniensis
MN7
D - Steinheim
Insectivora Mioechinus intermedius; Galerix socialis; Heterosoricinae;
Soricinae; ?Proscapanus sansaniensis
Chiroptera Vespertilionidae; Eptesicus campanensis; Chiroptera indet.
Lagomorpha Prolagus oeningensis; Lagopsis verus; Eurolagus fontannesi
84
Rodentia Spermophilinus bredai; Microdyromys complicatus
miocaenicus; Miodyromys aegercii; Myoglis meini;
Megacricetodon minor; Megacricetodon germanicus;
Democricetodon mutilus; Democricetodon gaillardi;
Eumyarion cf. latior; Anomalomys gaudryi
Carnivora Paralutra jaegeri; lschyrictis mustelinus; Martes cf. filholi;
Proputorius sp.; Trochotherium cyamoides; Trocharion
albanense; Amphicyon steinheimensis; Amphicyon sp.;
Amphicyonopsis? serus; Hemicyon gtiriachensis; Ursavus cf.
intermedius; Pseudaelurus quadridentatus; Pseudaelurus
lorteti; Sansanosmilus jourdani; Semigenetta sansaniensis
Perissodactyla Anchitherium aurelianense; Aceratherium simorrense;
Lartetotherium sansaniense; Lartetotherium steinheimense;
Phyllotillon fraasi
Proboscidea Gomphotherium angustidens
Artiodactyla Listriodon splendens; Conohyus simorensis; Albanohyus
pygmaeus; Micromeryx flourensianus; Euprox furcatus;
Heteroprox larteti; Palaeomeryx eminens; Dorcatherium
crassum
MN8
CH- Anwil
lnsectivora Galerix socialis; Lanthanotherium sansaniense; Plesiosorex
schaffneri; Dinosorex pachygnathus; Plesiodimylus chantrei;
Metacordylodon schlosseri; Alloscapanus sansaniensis;
Scaptonyx edwardsi; Talpa minuta; Desmanella stehlini
Chiroptera Scotophilus sp.; Tadarida sp.; Nyctinomus helveticus;
Miomegaderma gaillardi
Lagomorpha Prolagus oeningensis; Lagopsis verus; Eurolagus fontannesi
Rodentia Spermophillinus bredai; Albanensia albanensis; Miopetaurista
gaillardi; Forsythia gaudryi; Blackia miocaenica; Sciuropterus
sp.; Deperetomys hagni; Cricetodon sp.; Megacricetodon
schaubi; Megacricetodon similis; Megacricetodon germanicus;
Democricetodon brevis; Democricetodon freisingensis;
Eumyarion latior; Anomalomys gaudryi; Neocometes brunonis;
Microdyromys koenigswaldi; Microdyromys complicatus;
Paraglirulus werenfelsi; Paraglirulus conjunctus; Glirudinus
undosus; Eomuscardinus aff. sansaniensis; Muscardinus sp.;
Miodyromys aegercii; Miodyromys hamadryas; Paraglis sp.;
Myoglis meini; Keramidomys mohleri; Keramidomys
anwilensis; Eomyops catalaunicus; Trogontherium minutum
Carnivora Trochotherium cyamoides; Leptoplesictis filholi; Semigenetta
sansaniensis; Ursavus brevirhinus; Pseudarctos bavaricus;
Carnivora indet.
Perissodactyla Anchitherium aurelianense; Rhinocerotidae indet.
Artiodactyla Listriodon splendens; Conohyus simorrensis; Suidae indet.;
Albanohyus pygmaeus; Dorcatherium sp.; Micromeryx
flourensianus; Euprox furcatus; Cervidae indet.; Palaeomeryx
sp.; Eotragus sp.
(continued)
85
Table Z (continued)
MN9
E - Can Llobateres
Primates Rahonapithecus sabadellensis; Dryopithecus brancoi;
Sivapithecus sp.?
lnsectivora Plesiodimylus chantrei; Metacordylodon sp.; Postpalerinaceus
vireti; Parasorex socialis; Lanthanotherium sanmigueli; Talpa
minuta; Talpa vallesensis; Dinosorex aff. sansaniensis;
Miosorex grivensis; Crusafontina excultus
Chiroptera Chiroptera sp. 1; sp. Z
Lagomorpha Lagopsis cf. verus; Amphilagus fontannesi; Prolagus crusafonti
Rodentia Progonomys cathalai; Hispanomys thaleri; Megacricetodon cf.
minor; Eumyarion leemanni; Cricetulodon sabadellensis;
Anomalomys cf. gaillardi; Eomyops catalaunicus;
Keramidomys pertesunatoi; Heteroxerus grivensis;
Heteroxerus cf. rubricati; Spermophilinus bredai; Albanensia
grimmi; Miopetaurista crusafonti; Eomuscardinus vallesiensis;
Muscardinus crusafonti; Paraglirulus werenfelsi; Paraglirulus
conjunctus; Glirudinus undosus; Myoglis meini; Steneofiber
jaegeri; Steneofiber depereti; Trogontherium minutum
Carnivora Indarctos vireti; Ursavus primaevus; Ursavus brevirhinus;
Protursus simpsoni; Amphicyon major; Simamphicyon
batalleri; Canidae indet.; Martes mellibulla; Martes sp.;
Ischyrictis petteri; Sabadellictis crusafonti; Circamustela
dechaseauxi; Marcetia santigae; Mesomephitis medius;
Promephitis pristinidens; Trocharion albanense; Paralutra sp.;
Trochictis narcisoi; Semigenetta ripolli; Protictitherium
crassum; Protictitherium gaillardi; Hyaenidae indet.;
Machairodus aphanistus
Perissodactyla Hipparion catalaunicum; Chalicotherium grande; Tapirus
priscus; Dicerorhinus sansaniensis; Dicerorhinus
schleiermacheri; Aceratherium incisivum; Alicornops
simorrense
Proboscidea Prodeinotherium laevius; Deinotherium giganteum;
Tetralophodon longirostris
Hyracoides Pliohyrax sp.
Artiodactyla Albanohyus pygmaeus; Parachleuastochoerus crusafonti;
Listriodon splendens; Hyotherium soemmeringi; Micromeryx
flourensianus; Euprox dicranocerus; Dorcatherium sp.;
Capreolus sp.; Miotragocerus aff. panoniae
MN 10
E - Masia del Barbo (Teruel)
Insectivora Galerix sp.; Postpalerinaceus vireti; Crusafontina excultus
Lagomorpha Prolagus crusafonti
Rodentia Hispanomys peralensis; Hispanomys sp.; Democricetodon aff.
sulcatus; Kowalskia aff. fahlbuschi; Progonomys hispanicus;
Progonomys calathai; Muscardinus crusafonti; Tempestia
hartenbergeri; Eliomys sp.; Spermophilinus aff. bredai;
Heteroxerus; Atlantoxerus adroveri; Palaeomys castoro1des;
Dipoides problematicus
Carnivora Adcrocuta eximia
86
Perissodactyla Hipparion cf. primigeniurn; Aceratherium incisivum
Artiodactyla Decennatherium pachecoi
MN 11
E - Crevillente Z
Insectivora Galerix sp.; Dibolia sp.
Lagomorpha Prolagus crusafonti
Rodentia Kowalskia fahlbuschi; Parapodemus lugdunensis; Occitanomys
sondaari; Valerymys vireti; Eliomys truci; Muscardinus sp.;
Heteroxerus aff. grivensis
Carnivora Indarctos atticus; Mustelidae indet.; Plioviverrops guerini;
Amphimachairodus giganteus; Paramachairodus ~
Perissodactyla Hipparion sp.; Dicerorhinus schleiermacheri
Proboscidea Tetralophodon longirostris
Artiodactyla Microstonyx major; Dorcatherium naui; Eostylocerus aff.
pierensis; Cervidae indet.; Birgerbohlinia schaubi; Giraffinae
indet.; Tragoportax gaudryi
MNIZ
Los Mansuetos (E - Teruel)
Jnsectivora Galerix sp.; Dibolia major; Desmanella crusafonti; Talpa
minuta; Blarinella dubia; Paenelimnoecus repenningi
Chiroptera Megaderma vireti
Lagomorpha Prolagus crusafonti; Alilepus turolensis
Rodentia Ruscinomys schaubi; Parapodemus barbarae; Occitanomys
adroveri; Valerymys turoliensis; Spermophilinus turolensis;
Atlantoxerus adroveri; Eliomys cf. truci; Dipo1Cles
problematicus
Carnivora Canis cipio; Enhydriodon lluecai; Baranogale adroveri;
Adcrocuta eximia; Thalassictis adroveri; Plioviverrops guerini;
Lycyaena; Paramachairodus giganteus; Metailurus parvulus
Perissodactyla Hipparion concudense aguirrei; Aceratherium incisivum
Artiodactyla Euprox dicranocerus; Gazella deperdita; Tragocerus
amalthaeus; Hispanodorcas
Proboscidea Tetralophodon longirostris
UN 13
El Arquilla 1 (E- Teruel)
Insectivora Galerix sp.; Desmanella crusafonti; Blarinella dubia
Lagomorpha Prolagus crusafonti; Alilepus turolensis
Rodentia Ruscinomys schaubi; Cricetus cf. kormosi; Valerymys
turoliensis; Stephanomys ramblensis; Occitanomys adroveri;
Apodemus gudrunae; ?Myomimus sp.
Carnivora Sivaonyx lehmani; Adcrocuta eximia; Thalassictis adroveri;
Lycyaena sp.; Machairodus giganteus; Metailurus parvulus;
Fortunictis acerensis
87
Table Z (continued)
Perissodactyla Hipparion periafricanum; Hipparion primigenium truyolsi;

Hipparion gromovae gromovae; Dicerorhimus schleiermacheri
Proboscidea Zygolophodon turicensis
Artiodactyla Hexaprotodon crusafonti; Euprox sp.; Gazella deperdita;
Tragoportax sp.
MN 14
P - Podlesice
Insectivora Talpa minor; Desmana nehringi; Sorex alpinoides; Sorex
dehneli; Sorex runtonensis; Sorex minutus; Petenyia hungarica;
Paranourosorex ~ Blarinoides mariae; Petenyiella gracilis;
Soriculus kubinyii
Chiroptera Rhinolophus delphinensis; Rhinolophus grivensis; Miniopterus
schreibersi; Plecotus crassidens; Myotis podlesicensis; Myotis
danutae; Myotis dasycneme subtilis; Myotis cf. aemulus;
Myotis cf. exilis
Carnivora Baranogale helbingi; Vormela cf. petenyi; Mustela sp.
Rodentia Sciurus cf. warthae; Tamias cf. orlovi; Pliopetaurista cf.
dehneli; Blackia polonica; Pliopetes hungaricus; Pliopetaurista
sp.; cf. Sciurotamias; Leptodontomys aff. catalaunicus;
Keramidomys mohleri; Estramomys sp.; Protozapus
intermedius; Anomalomys sp.; Parapodemus coronensis;
Kowalskia magna; Kowalsia polonica; Cricetus sp. 1; Cricetus
sp. Z; Epimeriones progressus; Baranomys kowalskii; Prosomys
insuliferus; "Trilophomys" canterranensis; Glirulus pusillus;
Muscardinus pliocaenicus; Muscadinus cf. dacicus; Glis minor
Lagomorpha Ochotonid indet.; Leporid indet.
MN 15
F- 66 Perpignan (Serrat d'en Vacquer)
Primates Mesopithecus monspessulanus; Dolichopithecus ruscinensis
Insectivora Galerix depereti, Talpa sp.; Episoriculus gibberodon
Lagomorpha Prolagus depereti; Oryctolagus laynensis
Rodentia Mimomys davakosi-occitanus; Ruscinomys europaeus;
Trilophomys pyrenaicus; Trilophomys schaubi; Blancomys
neglectus; Cricetus angustidens; Stephanomys donnezani;
Castillomys crusafonti; Occitanomys montheleni; Paraethomys
meini; Paraethomys jaegeri; Rhagapodemus hautimagnensis;
Apodemus dominans; Apodemus jeanteti; Glis minor;
Muscardinus pliocaenicus; Eliomys intermedius; Pliopetaurista
pliocaenica; Castor praefiber; Hystrix primigenia
Carnivora Dinofelis diastemata; Felis issiodorensis; Felis aff.
maniculata; Viverra pepraxi; Pachycrocuta pyrenaica;
Nyctereutes donnezani; Canis michauxi; Canis adoxus; Ursus
ruscinensis
Perissodactyla Hipparion crassum; Dicerorhinus megarhinus; Dicerorhinus
miguelcrusafonti; Tapirus arvernensis minor
Proboscidea Anancus arvernensis; Zygolophodon borsoni
Artiodactyla Korynochoerus provincialis; Sus arvernensis minor;
Croizetoceros pyrenaicus; Paracervulus australis; Gazella aff.
borbonica; Alephis ~
88
Tubulidentata Orycteropus deperetl
Sirenia Felsinotherium
MN 16
I- Villafranca d'Asti (Triversa)
Primates Macaca florentina
Rodentia Castor sp.
Carnivora Viverra cf. pepratxi; Acinonyx pardinensis; Chasmaporthetes
lunensis
Perissodactyla Dicerorhinus jeanvireti; Tapirus arvernensis
Proboscidea Anancus arvernensis; Zygolophodon borsoni
Artiodactyla Sus arvernensis minor; Cervus cf. cusanus; Leptobos
stenometopon
MN 16
I- Villafranca d'Asti (Arondelli)
Insectivora Blarinotdes mariae; Episoriculus gibberodon; Sorex sp.;
Petenyia hungarica; Beremendia fissidens; Talpa cf. minor
Lagomorpha Prolagus savagei; Hypolagus sp.
Rodentia Mimomys polonicus; Mimomys (Cseria) gracilis; Apodemus cf.
alsomyoides; Muscardinus; Glirulus pusillus
Carnivora Parailurus cf. hungaricus; Baranogale helbingi
MN17
F- 26 Saint Vallier
Primates Macaca cf. florentina
Lagomorpha Ochotonid indet.; Oryctolagus lacosti
Rodentia Mimomys pliocaenicus; Hystrix refossa; Castor plicidens
Carnivora Nyctereutes megamastoides; Vulpes alopecoides; Felis (Lynx)
issiodorensis; Panthera schaubi; Acinonys pardinensis;
Megantereon megantereon; Homotherium crenatidens;
Pachycrocuta perrieri; Chasmaporthetes lunensis; Enhydrictis
ardea; Baranogale helbingi; Aonyx bravardij Meles thorali;
Ursus etruscus
Perissodatyla Equus stenonis vireti; Dicerorhinus etruscus
Proboscidea Anancus arvernensis; Mammuthus meridionalis
Artiodactyla Croizetoceros ramosus medius; Cervus philisi valliensis;
Euctenoceros senezensis; Leptobos stenometopon; Gallogoral
meneghini; Gazella borbonica; Gazellospira torticornis
89
The Astaracian is an age with less generic changes, There are difficulties to
delimit the MN 7 unit with generic events; these difficulties occur partially with the
classical placement of the Manchones fauna in MN 6. In fact, this local fauna is more
young than the Sansan local fauna. The Caproid~s migration (visible in the Manchones
fauna) is an important generic event as shown by Koehler (1988).
Some events, formerly associated with the early Turolian, such as the occur-
rence of several Hipparion or the first appearance of Parapodemus, seem now to occur
in late Vallesian. The Austrian locality Kohfidisch fits better in late Vallesian than
early Turolian.
The subdivision of Turolian into three units is relatively easy in western Europe
but more difficult in eastern Europe and west Asia. For this reason the position of
eastern localities are only suggested here without absolute certainty. The first
appearance of Hystrix seems to occur earlier in North Africa and eastern Europe than
it does in western Europe.
The Ruscinian has few new records other than the discovery of Mimomys in
Perpignan (Serrat-d'en-Vacquer), which is a good indicator for MN 15.
Persons studying "Villafranchian" faunas generally work with precise biological

events. It seems that the arrival of many important Villafranchian genera occur not
exactly at the same time and in western Europe will allow the further subdivision of
MN zones.
During the Miinchen Symposium in 1975 about mammalian stratigraphy

(Fahlbusch, 1976), some localities were selected to illustrate each MNzone. It.seemed
useful to give here (table Z) the complete mammalian faunal list of these localities.
REFERENCES
Fahlbusch, V., 1976. Report on the Internatioanl Symposium on Mammalian Stratig-

raphy of the European Tertiary. Newsletter Stratigraphy, v. 5 (Z/3), p. 160-167.
Kohler, M., 1988. Boviden des tUrkischen Miozans (Kanozoikum und Braukohlen der
Tiirkei. 28). Paleont. i Evol. t. 21, 1987, p. 133-246.
Mein, P., 1975. Resultats du groupe de travail des vertebres: Biozonation du Neogene
mediterraneen a partir des mammiferes, in Senes, J. (ed.), "Report on Activity
of the RCMNS Working Groups (1971-1975f," Bratislava, p. 78-81.
a
Mein, P., 1976. Biozonation du Neogene mediterraneen partir des mammiferes, in
Proceedings, VIth Congress, RCMNS, Bratislava, September 4-7, v. z.
90
MUROID RODENT BIOCHRONOLOGY OF THE NEOGENE
AND QUATERNARY IN EUROPE
Oldrich Fejfar
Geological Survey Prague

Malostranske n. 19
118 21 Praha 1, Czechoslovakia
Wolf-Dieter Heinrich
Museum of Natural History of the Humboldt-University

Invalidenstr. 43
1040 Berlin, German Democratic Republic
INTRODUCTION
The mammalian biochronology of the continental Neogene and Quaternary in

Europe has made remarkable advances. The present state of knowledge was discussed
at the symposium on "European Neogene Mammal Chronology" at Schlo~ Reisensburg,
May 1988. Investigations of fossil rodents have yielded important contributions to the
biochronology of the late Cenozoic in Europe. This is due to the rapid evolution of
many rodents providing short-ranged species that become characteristic index
fossils. In addition, the analysis of evolutionary trends within several rodent lineages
has yielded important biochronologic time markers such as the Promimomys-
Mimomys-Arvicola lineage.
The features outlined below enable us to utilize a biochronologic framework for

the European late Cenozoic. The authors of the present paper have presented several
contributions to that topic (cf. Fejfar and Heinrich, 1981, 1987). Our biochronologic
framework is based on the classical biochronological principle. It means the conclu-
sions result without exception from the chronologie ranges of species and genera. This
procedure provides distinct delimitations of faunal units. It is well known that the
mammal ages of the continental late Cenozoic in Europe (e.g., Ruscinian, Villanyian,
Biharian) have been insufficiently defined. More recent discoveries have shown that
these faunal units can be defined substantially better on the basis of concurrent ranges
of muroid genera (figure 1).
Nevertheless, the biochronologic conclusions derived from the concept of con-

current ranges of muroid genera have not been generally accepted. Objections were
discussed at the symposium "Evolution, Phylogeny and Biostratigraphy of Arvicolids
(Rodentia: Mammalia)" held at Rohanov (Czechoslovakia) in 1987 and during the
Reisensburg symposium on "European Neogene Mammal Chronology" in 1988. These
objections concern in the first line the number and names of the mammal ages distin-
guished. Thus, for instance, it was recommended to remove the term Villafranchian
from our biochronologic framework because that name is widely used in various senses
by different authors. Moreover, the emendation of the Ruscinian along with the
establishment of the Montpellerian (cf. figure 1; for details see Fejfar and Heinrich,

Edited by E. H. Lindsay el at.
Chronostratigraphy Continental Biostratigraphy
a in CE>ntral and Western Europe
:;[
Fejtar & Heinrich, 1980,,981,1987,1988
-~
QJ Global Mediterran MD Mammal Rodent Rodent MD Mammal Rodent Rodent
E /
i= MN Ages Stages Zones ~ Ages Stages Zones
:>, Holocene
Ty...u.b_e...Q.iQ!l MD Toringian
Arvico\o A. terrestris MD Toringian
Arvicola A. terrestris
2 Microtus A.cantiona Microtus A.conliano
L.. Mitazzian
0,5 a f-o,-,---
cL.. Stctlian
P\eis tocene ~~ilia~=- MQ Microtus MD Microtus M imomys
1 2 1 Bihar ian Bihar ian
a Colabrian 1 Mimomys 1 Mimomys savini
::J
0
2 MN Logurodon MN Lagurodon Mimomys

2 Vil!anyian Villanyian
PiacE'n- 17 Villanyia 17 Vil\anyia pliocaenicus
cion MN Mimomys MN Mime mys
Villa- Borsodia Vitl a- Borsodia polonicus
'3 ~
16 b potonicus
3 ~
MN Mimomys
K f ranchian Dolomys Mimomys fran chien Dolomys
16a hajnackensis 16 a hajnackensis
MN Mimomys MN Mimomys
\~ MN
Pliocene Csarnotan
15 b Promimomys 15 b Mimomys occi ta nus
Ruscinian
4
Zanclian \ 15a
Ruse inion Promrmomys
"MN
15 a
Ruse inomys
Mimomys
davakosi
~ MN
MN Promimomys
14 b ~ Mont- Promimomys moldavicus
5 5 M'N MN pellierian Ruscinomys Promimomys
14 a 14a insuliferus
MN MN Stephanomys
iZ
Messinian
6 :>, 13 13 ramblensis
L..
a I-- f--
-
·-
\
L..
7 QJ MN MN Valerymys Parapodemus
1- Turolian
12 12 Hispanomys gaudryi
8 8 - -
MiocE>ne
~
Tortonian
MN MN Parapodemus
11 11 l ugdunensis
9
1---
MN MN Progonomys Progonomys
10 10 Rotundomys hisponicus
10 10 I--
1--- Valles ian
MN MN
Serravallian 9 9
lu.part)
11
Fig. 1. Chronostratigraphy and changing framework of muroid rodent bio-
chronology of the late Neogene and Quaternary in central and
western Europe, showing problems in the subdivision and nomination
of biochronologic units. For details see text.
1987, 1988) have not been accepted because this procedure would contradict the
conclusions of the 1975 Munich symposium on mammalian biostratigraphy of the
European Tertiary.
There is as yet no clear decision to this discussion; however, we wish to raise the
point that our basic biochronologic concept is not at issue with these objections. Our
previous proposals, with some minor deviations from the Munich recommendations,
can be easily adapted without any substantial consequences for the biochronologic
concept that we have adopted. Contrary to other proposals, the boundaries of the late
Neogene and Quaternary mammal ages are clearly defined by the concurrent ranges of
rodent genera. In this sense, the establishment of both the Ruscinian sensu stricto, as
well as the Montpellerian, remains open for further discussions in thesa:Die way that
Ruscinian may be applied for the time span of the MN zones 14 and 15 in the sense of
the Munich recommendations. In order to facilitate an agreement between different
92
Revised Continental Chronostratigraphy
0
Biostratigraphy Ranges of significant L
in Central and Western Europe c
rodent genera Q)
MQ Rodent Mediterran Global
Mammal Rodent in Central and Western Europe E
/MN Ages Super zones Zones t=
I
Arvic.ola A.terrestris Ty_rrh~ian >.
MQ Toringian Holocene/
2 Microtus A.cantiana Millazzian 0c 0,5
Mimomys siCiliOn-- ._
MQ Microtus r:.;:v~i __
Mimor:nys
--------- ---- ~ 1 E:ffiWQP-=--=- Pleistocene Q)
~ 1
Biharian SOVInl - ·;;: 0
1 Mimomys Calabrian
Mimomys ~ ::J
pusillus a
MN Mimomys
2 2
17 pliocaenicus ~
Piacen-
Borsodia ~- '-....
M"N Villanyian Mimomys c ~ cion
Villanyia polonicus .g -~
vrJ
~ e:>:-
MN Mimomys ~ 3
16 a ha·nackensis _§'
MN
1/t
Mimomys .Q.Q
Pliocene
~
occitanus "'"
E~-
MN 4
v
Mimomys l/4
t
~;;~
"E
~
Trilophomys davakosi ~ Zanclian

15a >- E
'MN Ruse inion
Ruscinomys Promimomys o
c0
E
"" -;;:;o
0
moldavicus >-0
~ 0. ~
5
MN Promimomys -~
I>
0 ~
E
5
14 a insuliferus
:>:
~ ~-
MN Stephanomys "E E" ~

E _g E' Messinian
~
13
s ramblensis
.E _9- .§ I} >. 6
._
r- - c
.,
0
e ·->::- L
Q_
0
·-
0 ':2,
g
~
._
l
"
~
~
.,.,~
u
"5 7 7
1/
MN Turo- Vale>rymys Porapodemus Q)
gaudryi -~ f-
12 l ian
:>:
Hispanomys ~
:>: 0:
0:
'~
- - g_.2_
., ~ ~ 8 8
2 -~ >. Miocene
~
~ " E
MN -;:" Parapodemus ·s 2 ~ Tortoni an
lugdunensis
~ ~ .20
0
11 w
Ql · - g
o::;:oc
MN ~ Progonomys
10
~ hispanicus
- Vall•-
sian 1-- Rotundomys
Progonomys 10
MN ~ Microtocricetus
9 ~ mottassicus Serravallian
11 (u.part)
points of view, a revised tentative biochronologic framework of the late Neogene and
Quaternary in Europe is given below. It covers the time span from the Vallesian up to
the Holocene.
FRAMEWORK: LIMITATIONS AND POSSIBn.ITIES
The biochronologic method applied is based on a synopsis of ranges of about 60

muroid rodent genera recorded from the late Cenozoic of Europe. Great importance
has been attached to evidences of evolutionary changes in lineages as well as dispersal
events, especially the first appearance data of the rodent genera considered.
In our framework, zones are the lowermost biochronologic units. They are
mainly based on total ranges of rodent species from lineages, and each of them repre-
93
sents a distinct evolutionary and chronological level. This can be exemplified by the
members of the so-called Promimomys-Mimomys-Arvicola line. Superzones and
mammal ages as major biochronologic units have been characterized by alternating
combinations of successive ranges of muroid rodent genera. Significant features of
such combinations are (1) the one time occurrence, (Z) the short time span (range), and
(3) the relatively wide distribution, covering large parts of the European continent
(Fejfar and Heinrich, 1981, 1983, 1987). The lower boundary of a superzone or
mammal age is defined by the first appearance datum (FAD) of a genus. The upper
boundary is given by the FAD of a succeeding genus, defining the beginning of the next
younger superzone or mammal age. Additional, but less certain, information is
available from the disappearance, especially from the extinction of rodent genera,
i.e., the last appearance datum (LAD).
As with other biochronologic frameworks proposed, the procedure applied here

offers not only possibilities for expansion but also limitations. We are aware of this
and, therefore, we consider our framework a starting point for further more detailed
work in the future. Limitations result in the first line from our present stage of
knowledge, especially from gaps in the fossil record. There is, for instance, a very
limited number of Vallesian and Turolian sites in central and eastern Europe that have
provided remains of muroid rodents (figure Z). Due to the lack of fossils, the precise
chronologie range of several rodent genera has remained far from clear up to now.
Moreover, from the methodological point of view, special consideration must be given
to the possibility that FADs may not be strictly synchronous within larger regions such
as the European continent. Dispersal implies time. Therefore, the evaluation of FADs
must take into consideration regional retardation effects in the migration caused by
the capability and competition of the animals involved on the one hand, as well as the
paleogeographical and paleoecological accessibility on the other hand. FADs in
muroid rodent dispersal must be treated with caution as can be shown by the record of
arvicolid genera Synaptomys, Lemmus, Villanyia, and Borsodia. These genera
appeared in eastern Europe distinctly earlier (e.g., late Ruscinian) than in central
Europe (Villanyian), and some of them have never reached western Europe. This
clearly demonstrates that dispersal in muroid rodents can be a "time-transgressive"
process, occasionally involving geological time dimensions. On the other hand, the
well known Microtus (Allophaiomys) dispersal event has been demonstrated nearly
synchronous within Europe and Asia. This suggests tpat the utilization of FADs can be
reliable over great distance, and one should not be reluctant to try this application.
Further problems arise from the dispersal pattern of muroid ·rodents considered
in our framework. Some of these genera are recorded in wide geographical regions of
Europe, whereas others are distinctly endemic. Thus, for instance, recent discoveries
on the "index fossils" of the Ruscinian, Ruscinomys and Trilophomys, have suggested
that the distribution of Ruscinomys appears restricted to western and southern
Europe, whereas Trilophomys occurred from western to central Europe, including the
adjoining regions to the south. Until now, there has been no evidence that
Trilophomys had ever been endemic in eastern Europe. Consequently, the intraconti-
nental correlation of Ruscinian Trilophomys-Ruscinomys faunas with corresponding
faunal complexes of eastern Europe has to be supported with other guide fossils such
as Mimomys (Cseria) gracilis and Mimomys (Mimomys) occitanus, which were present
in western, southern, and eastern Europe as well as in adjacent regions. The muroid
rodents share this phenomenon with other Cenozoic mammals, and the problem out-
lined above demonstrates clearly the far-reaching significance of biochronologically
well defined time markers as well as regional biozonations that have been investigated
intensively.
Finally, we should like to raise another controversial subject: the taxonomic

identification of rodent taxa. Differences in taxonomic concept applied to muroid
rodents introduce considerable problems in the faunal correlation in the late Cenozoic
of Europe. This can be exemplified by Microtus, a genus widely used for the biozona-
tion of Biharian mammal age. Thus, for instance, a highly sophisticated classification
of Microtus in Austria (Rabeder, 1981, 1986) and France (e.g., Chaline, 197Z) is con-
fronted with a more conservative taxonomic concept used in other parts of Europe,
94
Significant localities
~ Chronostratigraphy Revised Continental
~ Biostratigraphy
~ ..... ~ _ in Central and Western Europe France Middle and South East Europe
Spain
England Europe Near East
Lezetxiki. Cueva del Agua. Ambrona ~:~g~~~~~r.:~7~~~o~~de. Toto. Buhlen. Touboch Godjo~S-kurlalov.Citremoinik1
El Hoguer6n. CuUar de Bozo 1.2 St Esteve-Jonson. Hoxn!l Hundsheim.Petersbuch.Mosbach Stretica,Cigirin,Gunki.Lichvin1
lama Ouemodo 1,2, HU11scar 2.3. ei~s~~~1~~u~~i~f~f~f.~ 1 - ~~11_~,~l;'~~!l~~~~:t::". P.tg:t!"n°:vkk6·. ~tr:k~~li~~t~~~

Los Yedros. PuE>rlo Lobo1.4, Bogur 2,
Coi'iodo de Murca 1,2, Vento M•cqno 1.2; Courtl!'rolles, Saont- Bt>tfoa 2. 5, 7,9, Mtiningen, 1.2,_1<os_lovo Balko .Novo-
Fuentenuevo 2,c,Orcq4-7,P, Sauveur. Mas Ramboult. VCet6re 1.4A/7. 38/1- trooekoJe Platovo 2 .I<Of'!Ji-
Barranco LeOn 1-3
~~~~~-~;~~l~g~~~/bn.y 1' Ooo7?~:1f~~1~FeJ1~:~~.~\e,'- g~~i~:l:tekho~~\~j~:ry,
MN Momomys Barranco de los Cosejos, Orce 2
Borsodia p\iocot>nocus ~i!~~J:,~~~~~~~~)~i~~ ~~r~~.~Bt~:!~r~r!~:~,ja:~Jgv~e:. Musoid. Liveneovko 1.2J,
2 Piacen- 17 Vi\lanyian ~~~~t~~~~~ ,r- 12:
Caiiodo Real. Jtes Medos Monlo9ny. Roca Ntyro. dorf,.Gundersheim 2, l<rems(zf Svopo
Viltanyia Chagny1, Montoussli 5 Mota Joskynoo 2. Solnhofom
cian
Mim polonocus Geo 1 ,Orct> D.C, Alquerio. Les Etouoires. Chogny 21 Montopoli, Za\11kovo Dolno, I KuSkuno. Livencovko 4.
Romanian
~ Concud -Villa9e. Fuentenuevo 1, Beoun 1. cessey -s -T, Remb1el1ce Krolewskoe. Navajo Etulijo 2. Uryv 2
Mom hojnack•nsis Concud -Siotoon, Bozo Mogny -les-Auxonnes Osztromos 7
16
3 MN Comenoil\es.Seynt>s, Arondelli-Tr•verso, Lovencovko S.l<otlovino.
~~~g,:;~~;~od~ A7u~':;tJ{~2~ 1rf{~~~ 0{"3~4a Bolorvc 2, Voolt>ltt> San Giusto. Arci!le. Kryi:onovko. Uryv 1
15 5; Cofiodo de Murcia 3, Sorrion HOJn6tko,Stronundorl0
0: 1--------l
N
Tri\ophomys
4
LJ2 b Ruscmian Ruscinornys
E~~~Jo~~~2~~cUge~~o~f~la2 ~:~8~ft>~:\.
Loyno . _
~f~e~outet. sete ?§}~~t;~e~~l~~~n:~o~~?~~ ·. ~~~;i~.~gV~~~~2~~i~Y·
w~ze. csornoto2
Zanet ian
~~f~!~f::·~~- 1 ~~~:~~~~~;~j~e.'l1:~o~:Cos. ~~~ ~.t>~r~~l·d~~~oc"qeune~- Ptolemois 3 ~~~~J~ .Rlo~~obond,
Pto\emo•s 1. Spolool.
5 Ce\odos 4o.~b.4c,8o,8b, Ald~huelo Vendorgues ~~~~·;,·, Mo\u~ten•
Celodos 3,g. La Jutoono, Piste de Cuevas, Houtomogne, Vollne_uve Dowronkhel 15,14,

MN PeroleJoS t>. Gorofe1 Terrots, Montpelloer I Podlesice,Moto Covet, Pul-t> Chorkhi,Antopovko
Ce\lneuve
La Albt>rco. Coravaco1 ,_Voltolbo Baja. Boccmetlo V3,

6 A_lcoy. Solabreilo, Crevollente 6, l<olilhies, Boltovolr, fitomoon. Amosyo,
""'-"' ~ lobrillo, La Fontana. Mosodo del Volle? Chol_koutso. Polgolrdi. CobruO:i.
Vale-rymys Voldect>bro 3, Vento del Mora. Mont so (Rh_J. Tiro spa\
Poropoderr<Js
~l Arquilla. Mo soda del Vo!te 6 Pikermo (Chomoteri J
Turolian
"'"I Hispanomys qoudryi
Pontian 12
7 r~:~~~.',·;,:~~·~,'·'·'·'' Lissieu. Rotovoux, Hot von, Formosovka, CimiSlio.
Concud 2.3.
Concvd C9, CC,CL, Stade de Cucuror1 (MI SUmeg, Toroklio. BoyirkOy.
MN Poropodemus ~I Crevillente 4,5, Lubo!ron).M•strolsi,li Bocconelto V2 Ghozghoy
lugdunensos Vil\otbo BaJO 2;
Tortojodo B.C, Aljezor B
8 MN Proqonomys 5~~~~08~:kheim. Bu~ory.

hisponocus Crevillente 1.2,3,
Vo"'"'"'· Per luis Kohlidisch, Kalla.
Tortonian Molton Boccinello V1
Alfombro, Vovero de Pinos Sherullah.
I-t" Tori.OJ. o. '" A.•. Los A.9uonoces, Cs6kv6r Cum ali
Biodrok [BootioJ Agha Jari Fm_ (lnjona),

5 Kostellios-Hill
9
Pannonian I
MN
9
Valles ian
Progonomys
Rotundomys
ERI
MicrolocritehJs
mollossicus
Torremarmo on 1,2
Mosio del gorbo 2A. 29,
I~~~~'J;~ 3~ 6~ H~j'or,
Fretroo de Roo Maoso,
Sob loy
Montredon
Aubi9nos
Suchomosty
Gotundorf
Vlis.,ndorf
Bayroktept l[.][
Kalla
10 Hommerschmiede,
~~::~~0\5~~J~~~{f~a~!~~~g~r~~~~· I Douf ~~~~e~t~~. I~~~0-i.s<~6\!gy,Mohmutk0y
Ampudio 8-80, 9; Torremormojon 5, HOweneg9. Eppelsheim
Serravallian Cubillos2
(u.port)
Fig. 2.. Chronological distribution of muroid rodent-bearing sites recognized in the late Neogene and
CD
U1 Quaternary of Europe.
such as Czechoslovakia (e.g., Fejfar and Horacek, 1983). Another striking example is
given by Borsodia and Villanyia which have been utilized for defining major faunal
units of the Pliocene in Europe. In accordance with other authors in central Europe
(e.g., Rabeder, 1981), we put the early rooted lagurids into the genus Borsodia (Fejfar
and Heinrich, 1983), a conclusion that is strongly supported by the enamel band differ-
entiation of the cheek teeth. But, this procedure runs contrary to a taxonomic con-
ception widely advocated in eastern Europe which assigns the early rooted lagurids to
the genus Villanyia (e.g., Aleksandrova, 1976; Topachevski and Skorik, 1977).
From all that, it can be concluded that we are far from attaining a comprehen-
sive biochronologic framework for all of continental Neogene and Quaternary fauna in
Europe. However, we are convinced this tentative biochronologic framework, based
on muroid rodents, is reasonable and should prove reliable. It should be treated as a
starting point for a comprehensive biochronologic framework that can be recognized
today only in its broadest outlines.
MUROID RODENT BIOZONATION OF THE LATE

NEOGENE AND QUATERNARY IN EUROPE
The chronological succession and subdivision of the muroid rodent superzones is

given in figure 3. The report is based on a large number of publications presented by
different authors, but not all of them can be quoted here. Their papers cover various
regions such as western and southern Europe (e.g., J.-P. Aguilar, J. Agusti, G.
Bartolomei, H. de Bruijn, J. Chaline, R. Daams, M. Freudenthal, C. de Giuli, Th. van
Kolfschoten, N. Lopez-Martinez, D. Mayhew, P. Mein, A. van der Meulen, A. Ruiz
Bustos, and A. van de Weerd); central Europe (e.g., D. Janossy, V. Fahlbusch, W. von
Koenigswald, L. Kordos, K. Kowalski, M. Kretzoi, A. Nadachowski, G. Rabeder, G.
Storch, and L. Thaler); and eastern Europe (e.g., K.A. Agadshanyan, L.P.
Aleksandrova, M.A. Erbajeva, A.K. Markova, A.N. Motuzko, C. Radulesco, P. Samson,
V.P. Suchov, E. Terzea, V.A. Topachevski, and V.S. Zazhigin). A selected bibliography
is given at the end of the present paper.
In the following review the type locality of a rodent superzone is considered

identical with the type locality of a mammal age insofar as the latter has yielded the
significant combination of rodent genera. The topic "significant taxa" considers
muroid and other rodent groups as well.
Progonomys-Rotundomys Superzone
Diagnosis: Temporal interval characterized by the concurrent ranges of Progonomys

and Rotundomys. The lower boundary is given by the FAD of Progonomys. The upper
boundary is indicated by the FAD of Valerymys.
Type locality: Can Ponsic (Marks, 1971; Aguirre et al., 1975; Van de Weerd, 1976).
Additional biochronologic data: The upper boundary of the Progonomys-Rotundomys

super zone is also characterized by the FAD of Collimys, Occitanomys, and
Epimeriones; the lower boundary by the FAD of Microtocricetus and Pliospalax. Most
likely the FAD of Hipparion coincides with the FAD of Progonomys.
Corresponding mammal age: Vallesian (Crusafont-Pairo, 1950).
Remarks: In central Europe the_ Vallesian mammal age is characterized by the per-
sistence of several Miocene rodent genera such as Democricetodon, Eumyarion,
Anomalomys, Myoglis, and Glirudinus. Most important is the occurrence of
ramapithecine and dryopithecine hominoids, as well as the first appearance of
Hipparion. In central Europe Microtocricetus molassicus and Democricetodon gaillardi
are of some significance.
96
Revised Continental
Selected Genera
.f ·£
Chronostratigraphy Biostratigraphy & c: ] ::: IIJ Ill -: Ill
., :] ~ ~ ~., ~ -~ ., ~ ~~~~ :o :Ill ~ "'~ :"' -~ ~ ~ ~ 111 E 111 ~
in Central and Western Europe
!e•~a;~~x-.:e;oe.c.:?~ ~~'~~ExS>-Ec;E>-~Gt>-_:111 -~E>.~~~~~E e=.~- >.~o :::~.,c "'
CIJO_~·~UEE O~o 0 :;-~.~e~>ootlcn~~U~~E~~CII:::IIII~o~E>.III~ 111 oEe~ 111 o.'!E'C~W~:J.e ::~ 111 c
~~!; ~-~~ ~ ~..;;~ ;~ ~ ~i: E ;_-~~g_.e~{~-; .=g ~=-~ ~.:~-~~g ~ E2 E~~ ~ E >.-~~~~·;:='go-;;~)(~ ~0
;= Stages MO Rodent ~ :~ E g,; &_ ::i ~~g'~ ~ ~ ;::~-~::3~ ~~ :~ ~ ~g g'-; ~~-~~~~~:; ~ ~2 ~ gg,~ ~2~ g ~~0 ~~ .;~~-; •::
Fig.3. Range chart of muroid ~ /
Mammal Rodent a. 111 111 ::1 01 c .I!!:=·- o ,_ o o > 111 o o a.u .. ·- 111 .... o 0 :J_,.c o= a. .. a.o .. ·;: o·---·;: o a.o- c ·-:::::: o > . 0 - • ,_ · - · - a.o o ._::I o
rodent genera and tentative • Ml<llt~rron
c~nlrol
Parat•thy& MN
Ages Superzones Zones zco~~<z~~~~~~m~u>wo~~-~~~~m~u<<uu~~m~~oc~~~~~>m~~u~~~c~w~c%~
Hat. Ty~~henial'! MO Toringion Arvicota A.t•rr..tr~s
Lllll!'
[IIIII
MiLiazzion 2 Microtus A.Uinliana
biochronologic subdivision of
011
~ Sicilian Mimomys
Microtus _S_!!.¥~1 __
l:: Emilian MO
the late Neogene and Quaternary 1
;; Biharian Mimomys :~'::1'!.'
'£ Calabrian
1 Mimamys
pu•mus
in central and west Europe. The
range chart is based on select- 2
MN
Borsodia
Mimomys
plklca.nicus
17
ed genera occurring in Europe Piacen-
cion
Villanyian
Villanyia
I
and adjoining areas. The
scheme is recognized such as 3
Romanian
~-;;;;;
16
lj;;;j
Ml"" polonicus
Mim.""""'k-~
rH
Ill
15
the very sporadic Ruse in ian 1--- 1-
records of Lemmus and 4 Zane lion
1--- 14b
MNI Ruse inion I TcHophomy•l
Ruscinomys
'""'-''
..,ldovocus
Synaptomys in eastern Europe
whose precise chronologie 5
oacion MN
14a
I r::~:::~:s
1
II
position needs further inves-
I I
~
tigations. Due to different
Stephanornys
MPssinian rambl•nsos
taxonomical evaluatings of
early lagurids, the eastern
6
1---
MNI Turolian
I Valerymys
Hispanomysl
ParapodPnOJs
~audryi
Ponti on 1_2
European findings of Borsodia
I ...........
llil
7 t-
and Villanyia assigned to
I
MN
lu9dunensi1
11
the late Ruscinian are not
considered here. Cronostrati-
8
Tortoni on 1---
MN
10 I I ~::::.~::
graphy according to F. Stein- t-
inger, Reisenburg Symposium, 9
I
1988.
10
I I~N~
Paman;an
Valles ion Progonomys
Rotundomys I
Mim:tocrUtus
fftOlklssicus
r---1
11 "'-ro.<allian
{u.parl)
""""'""" N Astaracim
....
CD
The Progonomys-Rotundomys superzone can be subdivided into two zones as
follows:
(a) Microtocricetus molassicus zone
Diagnosis: Temporal interval characterized by the range of Microtocricetus

molassicus to the range of Progonomys hi11panicus.
Type locality: Rudabanya (Kretzoi et al., 1974).
Remarks: Rudabanya has been elected as type locality because the site has yielded
both important floral and faunal assemblages comprising macro- and micro-mammals
as well; all of these taxa are inhabitants of a riparian forest belt.
Significant taxa: Microtocricetus molassicus, Democricetodon gaillardi, Anomalomys

gaillardi, Myoglis meini.
Localities: See figure z.

(b) Progonomys hispanicus zone
Diagnosis: Temporal interval characterized by the range of Progonomys hispanicus to

the range of Parapodemus lugdunensis (Van de Weerd, 1976).
Type locality: Masia del Barbo ZB (Van de Weerd, 1976).
Significant taxa: Progonomys hispanicus, Progonomys cathalai, Hispanomys

peralensis, Kowalskia fahlbuschi, Microtocricetus molassicus, Muscardinus crusafonti,
Tempestia hartenbergeri, Myomimus cf. dehmi, Eliomys truci, Eozapus intermedius,
Chalicomys jaegeri.

Remarks: In comparison with the previous zone the hypsodonty of Microtocricetus
has slightly increased.
Valerymys-Bispanomys Superzone
Diagnosis: Temporal interval characterized by the concurrent ranges of Valerymys

and Hispanomys. The lower boundary is given by the FAD of Valerymys. The upper
boundary is indicated by the FAD of Trilophomys.
Type locality: Los Mansuetos (Crusafont-Pairo, 1965; Aguirre et al., 1975; Van de
Weerd, 1976).
Additional biochronologic data: At the beginning of the Valerymys-Hispanomys

superzone the genera Collimys, Prospalax, Epimeriones, and Occitanomys appeared for
the first time in Europe.
Corresponding mammal age: Turolian (Crusafont-Pairo, 1965).
Remarks: The Turolian mammal age represents a time interval with late Miocene
relicts (Parapodemus lugdunensis zone) as well as an increasing number of murid taxa
successively more diversified (Parapodemus gaudryi zone and Stephanomys rambliensis
zone). The late Turolian (= Messinian) was a dry climatic phase. At that time a land
connection was established between North Africa and the Iberian Peninsula and other
parts of southern Europe as well.
According to Van de Weerd {1976), a subdivision of the Valerymys-Hispanomys

superzone is proposed as follows:
98
(a) Parapodemus lugdunensis zone
Diagnosis: Temporal interval characterized by the range of Parapodemus lugdunensis

to the range of Parapodemus gaudryi (Van de Weerd, 1976).
Type locality: Tortajada A (Van de Weerd, 1976).
Significant taxa: Parapodemus lugdunensis, Occitanomys sondaari, Valerymys vireti,

Hispanomys freudenthali, Kowalskia fahlbuschi, Eliomys truci, Eozapus intermedius.

(b) Parapodemus gaudryi zone
Definition: Temporal interval characterized by the range of Parapodemus gaudryi to

the range of Stephanomys rambliensis (Van de Weerd, 1976).
Type locality: Los Mansuetos (Van de Weerd, 1976).
Significant taxa: Parapodemus gaudryi, Valerymys turoliensis, Occitanomys adroveri,

Hispanomys freudenthali, Ruscinomys schaubi, Kowalskia fahlbuschi, Eliomys truci,
Muscardinus aff. crusafonti, Spermophilinus turoliensis, Atlantoxerus adroveri,
Dipoides problematicus.

(c) Stephanomys rambliensis zone
Definition: Temporal interval characterized by the range of Stephanomys rambliensis

(Van de Weerd, 1976).
Type locality: Valdecebro 3 (Van de Weerd, 1976).
Signficiant taxa: Parapodemus gaudryi, Valerymys turoliensis, Occitanomys adroveri,

Stephanomys rambliensis, Apodemus gudrunae, Ruscinomys schaubi, Kowalskia
fahlbuschi, Cricetus cf. kormosi, Eliomys truci, Spermophilinus turoliensis, Dipoides
problematicus.

Trilophomys-Ruscinomys Superzou.e
Diagnosis: Temporal interval characterized by the concurrent ranges of Trilophomys

and Ruscinomys. The lower boundary is defined by the FAD of Trilophomys. The
upper boundary is given by the FAD of Borsodia (central Europe) and the LAD of
Ruscinomys.
Type locality: Serrat d'en Vacquer (Mein and Aymar, 1984).
Additional biochronologic data: At the beginning of the Trilophomys-Ruscinomys

superzone the genera Celadensia, Promimomys, Baranomys, and Micromys appeared
for the first time. Moreover, in central Europe the upper boundary of the
Trilophomys-Ruscinomys superzone is indicated by the FAD of Borsodia and
Villanyia. In other parts of Europe the boundary between the Trilophomys-Ruscinomys
superzone and the Borsodia-Villanyia superzone is clearly defined by the FAD of
Mimomys (Mimomys) hajnackensis and Mimomys (Cseria) stehlini.
Corresponding mammal age: Ruscinian (Kretzoi, 196Z).
Remarks: We prefer this combination of rodent generic ranges because the nomi-
nated taxa permit correlations between central and western Europe. For the early
99
part of the Ruscinian mammal age (Promimomys insuliferus zone and Promimomys
moldavicus zone) the presence of Promimomys, the absence of Mimomys, and the
FAD's of Celadensia, Trilophomys, Baranomys, and Micromys are significant. The
younger part of this mammal age (Mimomys davakosi zone and Mimomys occitanus
zone) is characterized by the first appearance of cementless, mesodont, and rooted
representatives of the genus Mimomys.
More recent results of rodent biochronology in Europe have shown the replace-
ment of Promimomys by Mimomys. According to the fossil record (Van de Weerd,
1979), Mimomys evolved from Promimomys. Therefore, there is no further reason to
separate a distinct Promimomys-Mimomys stage (Csarnotan) mainly based on the
occurrence of Promimomys in the karst fissure site Csarn6ta Z (for details see Fejfar
and Heinrich, 1980, 1981, 1983). Moreover, the fossil record of Promimomys at
Csarnota Z is based on an unstratified surface find. The supposed occurrence of
Promimomys in the type locality at Serrat d'en Vacquer (Mein and Aymar, 1984) is also
not justified (Mein, oral comm.).
Considering the ranges of Promimomys and Mimomys species, the Ruscinian

mammal age (respectively, the Trilophomys-Ruscinomys superzone) can be subdivided
into four zones as follows:
(a) Promimomys insuliferus zone
Diagnosis: Temporal interval characterized by the total range of Promimomys

insuliferus (Fejfar and Heinrich, 1987).
Type locality: Podlesice (Kowalski, 1956; Agadshanyan and Kowalski, 1978).
Significant taxa: Stephanomys medius, Paraethomys anomalus, Apodemus primaevus,

Micromys paricioi, Ruscinomys europaeus, Kowalskia magna, Kowalskia polonica,
Celadensia nicolae, Epimeriones progressus, Cricetidae nov. gen. canterranensis.

(b) Promimomys moldavicus zone
Diagnosis: Temporal interval characterized by the total range of Promimomys

moldavicus (Fejfar and Heinrich, 1987).
Type locality: Malu~teni (Simionescu, 1980; Kormos, 193Z).
Significant taxa: Stephanomys margaritae, Rhagapodemus hautimagnensis,

Occitanomys brailloni, Apodemus dominans, Apodemus jeanteti, Ruscinomys
europaeus, Celadensia nicolae, Trilophomys castroi.

(c) Mimomys (Mimomys) davakosi zone
Diagnosis: Temporal interval characterized by the total range of Mimomys

(Mimomys) davakosi (Fejfar and Heinrich, 1987).
Type locality: Ptolemais 3 (Van de Weerd, 1979).
Significant taxa: Castillomys magnus, Castillomys crusafonti, Castillomys gracilis,

Orientalomys galaticus, Occitanomys brailloni, Apodemus dominans, Apodemus
jeanteti, Paraethomys meini, Stephanomys margaritae, Micromys kozaninensis,
Trilophomys castroi, Pseudomeriones abbreviatus, Kowalskia intermedia.
100
(d) Mimomys (Mimomys) occitanus zone

(Mimomys) occitanus (Fejfar and Heinrich, 1987).
Type locality: Sete (Thaler, 1966).
Remarks: A most important faunal break can be recognized at the beginning of the
Mimomys (Mimomys) occitanus zone. The first intensive dispersal event in the history
of arvicolids in the Holarctic region took place at that time. Moreover, the sudden
appearance of Dolomys, Germanomys, Stachomys, Ungaromys, Mimomys (Cseria), and
Pliomys reflects the first significant radiation of arvicolids.
Significant taxa: Castillomys magnus, Castillomys crusafonti, Castillomys gracilis,

Orientalomys galaticus, Occitanomys brailloni, Apodemus dominans, Apodemus
jeanteti, Paraethomys meini, Stephanomys margaritae, Kowalskia intermedia,
Trilophomys depereti, Trilophomys pyrenaicus, Mimomys (Cseria) gracilis, Propliomys
hungaricus, Dolomys milleri, Stachomys trilobodon, Germanomys helleri, Ungaromys
altenburgensis.

Remarks: The Csarnotan in the sense of Kretzoi (1959) corresponds to the Mimomys
(Mimomys) occitanus zone of the biochronologic framework presented here.
Borsodia-Villanyia Superzone
Diagnosis: Temporal interval characterized by the concurrent ranges of Borsodia and

Villanyia. The lower boundary of this super zone is given by the FAD of Borsodia, the
upper boundary by the FAD of Microtus (Allophaiomys) indicating the beginning of the
Biharian mammal age in the sense of Fejfar (1976) and Fejfar and Heinrich (1980,
1981, 1983).
Type locality: Villany 3 (Kretozoi, 1941; Janossy, 1986).
Corresponding mammal age: Villanyian (Kretzoi, 1941).
Remarks: The sites of Beremend (southern Hungary) assigned by Kretzoi (1956) and
Janossy (1986) to the early Villanyian (Beremendian) have provided two biochronolog-
ically important voles: Mimomys (Cseria) stehlini and Mimomys (Mimomys)
hajnackensis. By means of both species, a biozonation of the emended Villafranchian
(Borsodia-Dolomys stage in the sense of Fejfar and Heinrich, 1980, 1981, 1983) was
proposed by the present authors. In the meantime, however, it has turned out that the
term Villafranchian does not fit in the rodent biochronology. Moreover, there is no
reason to introduce a new name or to maintain the former Borsodia-Dolomys stage.
This faunal unit is abandoned on the strength of the following considerations.
The name Villanyian was introduced by Kretzoi (1941), but the upper and lower
limits of this faunal interval have never been precisely defined. Later, the Villanyian
was subdivided into two (Beremendian, Kislangian) or three (Beremendian, Tomanian,
Kislangian) smaller intervals (for details see ,Jcinossy, 1986). The new record of
Mimomys (Cseria) stehlini and Mimomys (Mimomys) hajnackensis in the Beremendian
type assemblage of the Beremend site, southern Hungary, indicates that the time span
of the Villanyian in the sense of Kretzoi (1941, 1956) includes the time span of the
Borsodia-Dolomys stage in the sense of Fejfar and Heinrich (1980, 1981, 1983). There-
fore, the latter unit is superfluous.
In comparison with other proposals, the framework of the Villanyian presented

here is distinctly defined and limited. The lower boundary of the Villanyian, the
Borsodia-Villanyia superzone, is set by the FADs of Borsodia and Villanyia in central
Europe. The upper boundary is given by the immigration event (FAD) of Microtus
101
(Allophaiomys). A biozonation of the Villanyian Borsodia-Villanyia superzone is pro-
posed as follows:
(a) Mimomys (Mimomys) hajnackensis zone

(Mimomys) hajnackensis (Fejfar and Heinrich, 1981).
Type locality: Haj!l~cka (Fejfar, 1964; Fejfar and Heinrich, 1985).
Remarks: At the beginning of the Mimomys (Mimomys) hajnackensis zone, cementum

appeared for the first time in the synclines of the molars of Mimomys. Simultane-
ously, the undulation of the enamel crown base increased. Moreover, the microstruc-
ture of the enamel in Mimomys displays further modifications (Fejfar and Heinrich,
1982).
Significant taxa: Mimomys (Cseria) stehlini, Mimomys (Mimomys) hajnackensis,

Mimomys (Kislangia) hintoni, Borsodia petenyi, Dolomys milleri, Germanomys helleri,
Stachomys trilobodon, Villanyia sp. (= Cseria opsia), Ungaromys sp., Estramomys
simplex, Prospalax priscus, Allocricetus ehiki, Apodemus alsomyoides.

(b) Mimomys (Mimomys) polonicus zone

(Mimomys) polonicus (Fejfar and Heinrich, 1981).
Type locality: R~bielice Krolewskie (Kowalski, 1960).
Remarks: Within the Mimomys (Mimomys) polonicus zone the lemmings (Synaptomys)
appear in central Europe. They are the first representatives of completely hypsodont
and rootless arvicolids in Europe.
Significant taxa: Mimomys (Cseria) stehlini, Mimomys (Mimomys) polonicus,

Ungaromys sp., Germanomys weileri, Stachomys trilobodon, Villanyia veterior,
Synaptomys (Synaptomys) europaeus, Dolomys cf. milleri, Estramomys simplex,
Pliopetaurista dehneli, Pliopetes hungaricus, Sminthozapus janossyi, Glis minor,
Dryomimus eliomyoides, Glirulus pusillus, Prospalax priscus, Baranomys loczyi.

(c) Mimomys (Mimomys) pliocaenicus zone

(Mimomys) pliocaenicus (Fejfar and Heinrich, 1981).
Type locality: Villany 3 (Kretzoi, 1956; Janossy, 1986).
Additional biochronologic data: Moreover, the Mimomys (Mimomys) pliocaenicus

zone can be distinctly characterized by the concurrent ranges of Lagurodon and
Villanyia. The total range of Mimomys (Cseria) reidi parallels that of Mimomys
(Mimomys) pliocaenicus.
Remarks: The origin of Mimomys (Mimomys) pliocaenicus coincides temporally with

the FAD of La~odon in central Europe. With La~urodon arankae and Lagurodon
praepannonicus Te hypsodont and rootless arvicohds appeared again in central
Europe. The enamel crown base and the hypsodonty increased independently in differ-
ent Mimomys lineages. Clethrionomys and Lemmus entered the fossil record in
central Europe for the first time.
102
Significant taxa: Mimomys (Cseria) reidi, Mimomys (Cseria) pitymyoides, Mimomys
(Cseria) tornensis, Mimomys (Kislangia) rex, Ungaromys nanus, Lemmus aff. lemmus,
Villanyia exilis, Borsodia hungarica, Lagurodon arankae, Lagurodon praepannonicus,
Estramomys simplex, Dryomimus eliomyoides, Prospalax priscus, Apodemus sylvaticus,
Apodemus alsomyoides, Allocricetus ehiki, Clethrionomys sebaldi.

Miaotua-Mimomya Superzcme
Diagnosis: Temporal interval characterized by the concurrent ranges of Microtus and

Mimomys. Lower boundary equals the FAD of Microtus (Allophaiomys); upper bound-
ary equals the FAD of Arvicola.
Type locality: Vi~lany 8 (Kretzoi, 1956, 1961; VanderMeulen, 1973; Janossy, 1986).
Corresponding mammal age: Biharian (Kretzoi, 1941).
Remarks: A striking feature of the Biharian mammal age is the radiation of Microtus
(for details see Chaline, 197Z; Van der Meulen, 1973; Rabeder, 1981, 1986). More
likely the immigration of Microtus (Allophaiomys) coincides approximately with the
base of the Eburonian (Fejfar, 1976; Fejfar and Heinrich, 1983).
Contrary to other determinations (e.g., Kretzoi, 1941, 1956, 1961; Van der
Meulen, 1973; Rabeder, 1981; Janossy, 1986), the lower boundary of the Biharian is
defined here by the FAD of Microtus (Allophaiomys), representing one of the most
important turning points in Pliocene and Pleistocene faunal history (Fejfar, 1976;
Repenning and Fejfar, 1977; Fejfar and Heinrich, 1983). This unambiguously defined
biochronologic time marker can be widely used within the Holarctic Region.
Biozonations of the Biharian have been proposed by different authors such as

Kretzoi (1956, 1961), Chaline (197Z), Van der Meulen (1973), and Rabeder (1981).
Nevertheless, the results obtained in previous subdivision of the Biharian mammal age
have been faced with many problems requiring further investigations. They are due
partly to the uniformity of the Biharian mammalian faunas. Moreover, repeated
climatic oscillations which were connected with true inland glaciations during the late
Biharian (Elsterian) influenced the succession of mammal communities. A gradual
differentiation in "warm" and "cold" faunas corresponding with cold and warm periods
of the Biharian became apparent. A reasonable biozonation of the Biharian mammal
age, therefore, can be arranged only with "index fossils" that realize two precondi-
tions: rapid evolution and wide dispersal during both cold and warm periods. Biharian
muroid rodent taxa which are capable of realizing these preconditions are represented
by Microtus and Mimomys savini.
In Microtus, the calculation of so-called AIL-indices (Van der Meulen, 1973) on

the one hand, and the analysis of the frequency distribution pattern of morphotypes
(Rabeder, 1981, 1986; Fejfar and Hor~~ek, 1983) on the other hand may provide useful
data concerning the evolutionary level of the genus as well as the temporal position of
vole-bearing strata. Considering Mimomys savini, the disappearance of enamel islets,
the gradual reduction of roots, the increasing height of crown, and the state of the
enamel band enable us to assess the temporal position of Biharian mammalian
communities.
Nevertheless, we must take into consideration the incompleteness of the data

available. So, for instance, at most Biharian localities in Europe the AIL-indices of
Microtus have not yet been calculated. Moreover, the enormous variability of the
molar pattern in Microtus and its bearing for the biochronology needs further intensive
study because many details of the evolution of this vole-lineage have been far from
clear. Therefore, we prefer a provisional and simplified biozonation of the Biharian
based on the rather well known ranges of the last two Mimomys species as follows:
103
(a) Mimomys (Mimomys) savini-Mimomys (Cseria) pusillus zone
Diagnosis: Temporal interval characterized by the concurrent ranges of (the early)

Mimomys (Mimomys) savini and Mimomys (Cseria) pusillus.
Type locality: Villany 5 (Kretzoi, 1956; Van der Meulen, 1973; Janossy, 1986).
Significant taxa: Dicrostonyx antiquitatis, Lemmus aff. lemmus, Prolagurus

pannonicus~ .Lagurodon arankae, Pliomys eniscolfis, Pliom_ys lenki Clethrionomvs cf.
glareolus, Microtus (Allophaiomys) deuca1on,icrotus (All~mys) phocaeDicus,
Citellus primigenius, Prospalax priscus, Apodemus leptodus, Apodemus cf. sylvaticus,
Apodemus mystacinus, Allocricetus ehiki, Glis cf. sackdillingensis.

(b) Mimomys (Mimomys) savini zone
Diagnosis: Temporal interval characterized by the partial range zone of (the

advanced) Mimomys (Mimomys) savini, ranging from the disappearance of Mimomys
(Cseria) pusillus to the entry of Arvicola.
Remarks: The boundary between the designated Biharian zones needs further inves-
tigations because the disappearance of Mimomys (Cseria) pusillus provides an insuffi-
cient time marker.
Type locality: Voigtstedt (Kretzoi, 1965).
Signifi<:ant taxa: Dicrostonyx simplicior, Lemmus aff. lemmus, Prolagurus

pannomcus, Pliomys episcopalis, Pliomys lenki, Clethrionomys glareolus, Pitymys
arvalidens, Microtus ratticepoides, Microtus arvalinus, Neodon gregaloides, Neodon
hintoni, Neodon schmidtgeni, Citellus primigenius, Sicista praeloriger, Spalax advenus,
Apodemus cf. sylvaticus, Cricetus praeglacialis, Allocricetus bursae.

Arvicola-Microtus Superzone
Diagnosis: Temporal interval characterized by the concurrent ranges of Arvicola and

Microtus. Lower boundary equals the FAD of Arvicola.
Type locality: Burgtonna (Fejfar and Heinrich, 1980, 1981, 1983; Heinrich and
Janossy, 1978a,b).
Corresponding mammal age: Toringian (Fejfar and Heinrich, 1980).
Remarks: The beginning of this superzone is indicated by the first appearance of

Arvicola. The succession of alternating glacial and interglacial mammal assemblages
is considered to be an important feature of this time unit. The present dispersal of
muroid rodents came into being during the Toringian.
Biometrical analysis of an evolutionary trend within the Arvicola-lineage pro-

vides well defined time markers which have been found important for the biozonation
of the Toringian. Using the biometrical data obtained, the Arvicola-Microtus super-
zone can be subdivided into two zones as follows:
(a) Arvicola cantiana zone
Diagnosis: Temporal interval characterized by the total range of Arvicola cantiana,

having enamel quotients i>o100 (Heinrich, 198Z, 1987).
Type locality: Mosbach-Z (Schmidtgen, 1911; Von Koenigswald and Tobien, 1987).
104
Remarks: In central Europe Arvicola appeared for the first time during the Elsterian,
and in eastern Europe during the Elsterian or within the Holsteinian. In western
Europe Arvicola apparently made its first appearance in the late Cromerian
(Cromerian IV; Von Kolfschoten, 1988).
Significant taxa: Arvicola cantiana, Arvicola aff. sapidus, Lagurus transiens, Lagurus
lagurus, Neodon hintoni, Neodon gregaloides, Neodon schmidtgeni, Microtus
ratticepoides, Microtus nivaloides, Microtus arvalis, Microtus agrestis, Microtus
subterraneus, Microtus brecciensis, Microtus oeconomus, Microtus gregalis, Pliomys
lenki ultimus, Pitymys arvaloides, Clethrionomys glareolus, Lemmus lemmus,
Dicrostonyx "torquatus", Apodemus sylvaticus, Cricetus cricetus, Cricetus major,
Allocricetus bursae.
Localities: See figure 2.
(b) Arvicola terrestris zone
Diagnosis: Temporal interval characterized by the total range of Arvicola terrestris,

having enamel quotients <100 (Heinrich, 1982, 1987).
Type locality: Burgtonna 2 (Heinrich and Jcinossy, 1978b; Heinrich, 1982, 1987).
Remarks: The lower limit of the Arvicola terrestris zone coincides in central Europe
approximately with the Eemian/Weichselian boundary.
Significant taxa: Arvicola terrestris, Arvicola sapidus, Lagurus lagurus, Eolagurus

luteus, Microtus arvalis, Microtus agrestis, Microtus subterraneus, Microtus cabrerae,
Microtus duodecimcostatus, Microtus oeconomus, Microtus gregalis, Microtus nivalis,
Clethrionomys glareolus, Lemmus lemmus, Dicrostonyx "torguatus", Citellus
citelloides, Citellus citellus, Citellus (Colobotis) superciliosus, Cricetus cricetus,
Apodemus sylvaticus, Apodemus flavicollis, Apodemus agrarius, Glis glis, Eliomys
quercmus, Muscardinus avellanarius.
SOME ASPECTS OF MUROID RODENT FAUNAL EVOLUTION
In the following a brief review of muroid faunal evolution in Europe during the
last 11 Ma is given. The report is based mainly on stratigraphic ranges of about 60
late Neogene and Quaternary muroid rodent genera. The diagrammatic representation
of the ranges and its chronostratigraphical relationship is shown in figure 3. The fossil
record is fairly good in Europe, a series of faunal breaks can be recognized within the
time span specified here.
A first considerable faunal break is observed between the Astaracian and

Vallesian muroid rodent faunas. It is indicated by the appearance of Fahlbuschia,
Rotundomys, Progonomys, Microtocricetus, Hispanomys, Pliospalax, and Kowalskia.
Neocometes and Deperetomys became extinct before the beginning of Vallesian land
mammal age. Megacricetodon, Democricetodon, and Eumyarion, which became
extinct at the close of Vallesian, were important elements of the Astaracian rodent
faunas in Europe. Parapodemus made its appearance during the Vallesian. The fossil
record clearly suggests that the cricetids which predominated for a long time in
Miocene rodent faunas declined distinctly during the late Vallesian. They were gradu-
ally replaced by murids.
The Turolian land mammal age is characterized by a relatively constant predom-

inance of murids. True and aberrant cricetids are present, but they are not abun-
dant. At the transition from the Vallesian to the Turolian land mammal ages, the
genera Collimys, Valerymys, Prospalax, Epimeriones, and Occitanomys entered the
European fossil record for the first time. Collimys is apparently restricted to the
105
early Turolian. During the middle and late Turolian in eastern Europe the occurrence
of the aberrant cricetids Microtoscoptes, lschymomys, and Aratomys is noteworthy.
Records of Microtoscoptes of the same or older age in China and North Ameirca
suggest an immigration from Asia.
Fossil rodent material recovered from the Turolian land mammal age in westem
and southern Europe is rich enough to demonstrate considerable faunal differences
within the Turolian. Thus, for instance, a distinct faunal break can be recognized in
the late Turolian. There is no question that this event coincides with the beginning of
the Messinian stage. Genera of both African (e.g., Calomyscus, Dendromus,
Myocricetodon, Pelomys, and Protatera) and Asiatic origin (e.g., Cricetus,
Pannonicola, and Paraethomys) appeared in Europe for the first time in this faunal
wave. Some of these newcomers survived into the Ruscinian land mammal age (e.g.,
Protatera, Paraethomys, and Ruscinomys), the Villanyian land mammal age (e.g.,
Blancomys and Rhagapodemus), the Biharian land mammal age (e.g., Castillomys), and
the Toringian land mammal age (e.g., Allocricetus, Apodemus, and Cricetus). Several
rodent genera became extinct to the close of the Turolian, such as Byzantinia,
Hispanomys, Valerymys, Microtoscoptes, Aratomys, and Ischymomys. The Messinian
faunal break is considered one of the most important tuming points within the
European muroid rodent succession. It was caused by unique environmental conditions
in the Mediterranean region and adjoining areas (Moya-Sola et al., 1984; Steininger et
al., 1985) which are correlated with the opening of migration corridors.
At the beginning of the Ruscinian land mammal age a further wave of new
elements reached Europe from the east. Among them were Promimomys, Baranomys,
and Micromys. Trilophomys is now recorded in western and central Europe; its origin
is not presently clear.
The early Ruscinian rodent material is of special interest as it includes the first
true arvicolid of Europe, Promimomys. It records the beginning of the "age of
arvicolids" ranging in Europe from the early Ruscinian up to the present. Most likely
Microtodon, known from late Turolian of central Asia (e.g., Ertemte; Schlosser, 192.4;
Schaub, 1934; Fahlbusch et al., 1983), gave rise to Promimomys. According to
Repenning and Fejfar (1977) records of Promimomys in both Europe and North
America were similar in age and may indicate a nearly synchronous west and east
dispersal out of Asia during the earliest Ruscinian. The fossil record of Promimomys
is still insufficient, but it permits recognition of two alternating species (P, insuliferus
and R_. moldavicus) which might witness gradual endemic evolution. Contrary to
previous statements, there is no reliable evidence that Promimomys survived into the
late Ruscinian of Europe.
Three important dispersal events of arvicolids are recognized during the time
span following the early Ruscinian. They mark three distinct intervals in arvicolid
history which can be characterized as follows:
(1) The first radiation took place within the late Ruscinian and can be placed at
the beginning of the Mimomys (Mimomys) occitanus zone. The following arvicolid
genera entered the fossil record in Europe for the first time: Pliomys (Propliomys),
Dolomys, Germanomys, Stachomys, Mimomys (Cseria), Synaptomys, Lemmus
Villanyia, and Borsodia. They are commonly associated with Mimomys (Mimomysf
which persisted from the early Ruscinian. The abrupt and nearly synchronous appear-
ance of about nine arvicolid genera is due to a dispersal related to strong climatic
changes which opened the way for a rapid immigration from Asia. It should be
mentioned that the late Ruscinian records of Lemmus, Synaptomys, Villanyia, and
Borsodia are restricted to eastern Europe. In central Europe the genera Synaptomys,
Villanyia, and Borsodia made their appearance during the early Villanyian, and
Lemmus have not been recorded there before the late Villanyian.
(2.) A further important radiation of arvicolids is revealed by the appearance of

Lagurodon and Clethrionomys. It seems likely that the earliest records of both genera
originate from the earliest part of the late Villanyian, that is the beginning of
106
Mimomys (Mimomys) pliocaenicus zone. At that time Lemmus is seen for the first
time in central Europe.
(3) The third important radiation of arvicolids is the appearance of the rootless
arvicolid genera Dicrostonyx, Prolagurus, and Microtus (Allophaiomys), suggesting a
new invasion of hypsodont arvicolids from Asia at the beginning of the Biharian.
Moreover, Spalax appeared in central Europe at that time. During the late Biharian
the steppe lemmings, Eolagurus and Lagurus, made their appearance in eastern
Europe. Lagurus entered central and western Europe at the beginning of the Toringian
and was repeatedly recorded from there. Eolagurus dispersed from central Asia to
Rumania and Greece, but it has not been recorded further west. It disappeared from
eastern Europe during Holocene times.
Mimomys (Cseria) became extinct in the course of the Biharian land mammal
age and the long-ranging taxon Mimomys (Mimomys) disappeared at the close of
Biharian and gave rise to Arvicola. After the Bihar1an the rootless voles with ever-
growing cheek teeth became the ruling arvicolids in Europe.
Pliomys (Pliomys) disappeared from central Europe during the early Toringian
(Holsteinian), but it may have persisted in western Europe into the late Toringian
(Weichselian). Arvicola, an index fossil of the youngest European mammal age, the
Toringian, appeared in Europe during the late Cromerian about 0.5 Ma before present.
Not much is known about the Toringian history of Mus and Rattus which belong
to the latest radiation of muroid rodents in Europe. According to records from
Rumanian (Rattus) and Hungarian sites (Mus), both genera apparently appeared in the
early Toringian (Holsteinian), but they did not become common until Holocene times.
Mus and Rattus are now widespread in Europe; they were introduced into several parts
of Europe by man in late Holocene times. The present distribution pattern of muroid
rodents was established during the late Toringian.
SOME ASPECTS OF MUROID RODENT MOLAR EVOLUTION
During the last 11 Ma the muroid rodents underwent a rapid evolution. Especial-
ly the cricetids and murids show an enormous diversification correlated with an
immense variability of the molar pattern and a high degree of parallelism in several
stocks. Various important molar patterns and their chronological relationships are
shown in figure 4.
During the Astaracian and the Vallesian land mammal ages, the first cricetids
with mesodont-lophodont molars appeared independently from different lineages such
as Microtocricetus molassicus, Democricetodon gaillardi, Rotundomys bressanus, and
the gerbillids. This is an important molar pattern which was also common during the
Turolian. Toward the close of the Turolian the cricetids with mesodont-lophodont
molars became distinctly more abundant. They include representatives of different
provenance such as invaders from the east (Microtoscoptes, Ischymomys, and
Aratomys) as well as advanced members of endemic lineages such as Hispanomys, a
genus that became extinct at the close of the Turolian.
Special consideration needs to be given to some aberrant "vole-like" cricetids

which were distributed in the eastern parts of Europe during the middle and late
Turolian: Microtoscoptes sp. and Ischymomys ponticus (Topachevski et al., 1978).
Thus, for instance, the "vole-like" Microtoscoptes precedes to a certain extent the
arvicolids in developing hypsodont-prismatic (microtoid) cheek-teeth with well differ-
entiated enamel triangles, a condition that was not realized by true arvicolids prior to
the late Ruscinian, more than 1 Ma later.
During the Ruscinian land mammal age, cricetine rodents with a conservative
brachyodont-lophodont dentition (e.g., Cricetus) as well as some with an advanced
mesodont-lophodont molar pattern (e.g., Baranomys) were widely distributed. More-
107
21 22 13 14
I ' " 26
f [Jfj 27
10
11 10 9 8 7 6
Vo lles i on Tu ro l i an
Ptogonomys • Rotundomys Voterymys • l-l•spanomys
MN 9 MN10 MN11 MN 13
Fig. 4. Review of the significant taxa of muroid rodents in the last 11.5 Ma, during
the mammalian ages Vallesian, Turolian, Ruscinian, Villanyian, Biharian, and
Toringian. The taxa are represented by occlusal views of the first lower
molar reduced to the same unified length. Some examples are presented in
labial views to show the gradually increasing hypsodonty.
1-ZO: Significant taxa of Vallesian (Progonomys-Rotundomys superzone): 1,Z,

Hispanomys nombrevillae (Can Llobateres, Nombrevilla); 3, Hispanomys aragonensis
(Pedregueras); 4, Hispanomys peralensis (Peralejos D); 5, Eumyarion leemani
(Rudabanya); 6, Democricetodon gaillardi (Rudabanya); 7,8, Kowalskia cf. fahlbuschi
(Suchomasty, Csakvar); 9, Democricetodon sulcatus (Pedregueras n C); 10,
Democricetodon nov. sp. (Suchomasty); 11, Rotundomys bressanus (Montredon); lZ,
Anomalomys gaillardi (Rudabanya); 13, Rotundomys sabadellensis (Can Llobateres); 14,
Pseudomeriones nov. sp. (Suchomasty); 15, Microtocricetus molassicus
(Hammerschmiede); 16, Megacricetodon ibericus (Nombrevilla); 17, Democricetodon
freisingensis (Hammerschmiede); 18, Collimys primus (Eichkogel); 19, Progonomys
cathalai (Montredon); ZO, Progonomys hispanicus (Peralejos C).
108
it
Tl
"
l~l
!!ll " "" .
42 43 18 79 80
41
••
111111 I~~
iii~ ~,, : : :
sa 59
5 4
Ruse inio n Vill a nyian B iharian oringian

AtviC:OIO -
l rilophOtnys. Auscinomys. Bor s.od io • Vi Uonyio
MIUOi uS
• M N 14 b • MN 15 b • MN16 • MN17 MO 1 M02
21-38: Significant taxa of the Turolian {Valerymys-Hispanomys super-

zone}: 21,22, Hispanomys freudenthali (Tortajada A, Masada del Valle 2}; 23,
Ruscinomys schaubi {Los Mansuetos); 24, Byzantinia pikermiensis {Pikermi); 25,26,
Valerymys turoliensis {Los Mansuetos}; 27, Ischymomys guadriradicatus {Ischim-
Petropavlovsk); 28, Valerymys vireti {Tortajada A); 29, Occitanomys adroveri {Aljezar
B); 30, Microtoscoptes praetermissus {Ertemte}; 31, Parapodemus lugdunensis
{Tortajada A); 32, Parapodemus barbarae (Los Mansuetos); 33, Apodemus gudrunae (La
Tour); 34, Stephanomys rambliensis (Valdecebro 3); 35, Occitanomys sondaari
{Tortajada A); 36, Pros alax petteri (Eichkogel); 37, Paraethomys jaegeri (Gorafe 2);
38, Pliospalax sotirisi Maritsa).
39-61: Significant taxa of the Ruscinian {Trilophomys-Ruscinomys
superzone): 39,46, Promimomys insuliferus {Podlesice); 40,47, Mimomys (Mimomys)
davakosi {Ptolemais 3); 41,44, Dolomys nehringi {Csarn6ta 2}; 42, Germanomys helleri
{Ivanovce A}; 43, Stachomys trilobodon {W\!ze); 45, Pliomys hungaricus {Csarn6ta 2);
{Figure explanation continued on next page.)
109
over, the first true arvicolids with mesodont-prismatic molars (e.g., Promimomys) are
recognized at that time. Promimomys probably developed from Microtodon which has
been reported from late Turolian deposits in Mongolia (Schlosser, 19Z4; Schaub, 1934;
Fahlbusch et al., 1983). Although the molar pattern of Promimomys differs little from
that of Microtodon and other "vole-like" Ruscinian cricetids, its jaw musculature
shows clearly the arvicolid condition (Repenning, 1968; Repenning and Fejfar, 1977).
Arvicolids are characterized by rooted or rootless prismatic cheek teeth having

a varying number of salient angles. A remarkable dental diversity can be observed
(figure 5). The first arvicolids are characterized by rooted cementless and prismatic
molars that are mesodont (e.g., Promimomys) or hypsodont (e.g., Mimomys davakosi,
Dolomys, and Pliomys). This molar pattern was modified by the acquisition of crown
cementum deposited between the enamel triangles. Further steps of evolution pro-
duced rootless, hypsodont and prismatic molars that are cemented (e.g., Lemmus,
Synaptomys, Microtus, and Arvicola) or cementless (e.g., Prolagurus, LaguruiJ,
Eola~, and Dicrostonix) and ever-growing. After the Biharian land mammal age
arviCOiS with rootless, ypsodont, ever-growing and prismatic molars have become
predominant.
Gradual evolutionary change in the molar pattern is observed in several lineages

of arvicolids. This can be exemplified by the Promimomys-Mimomys-Arvic ola lineage,
covering a time span of more than 5 Ma. The following dental morphological changes
in the first lower molar can be recognized (e.g., Chaline, 1987; Fejfar and Heinrich,
1983; von Koenigswald, 1983; Repenning and Fejfar, 1977; see figure 5): (1) increasing
hypsodonty, (Z) increasing size, (3) appearance of crown cement, (4) increasing undula-
tion of the enamel crown base, (5) disappearance of enamel islets, (6) reduction of
roots, and (7) change from the Mimomys-like to the Microtus-like enamel band differ-
entiation. The rate of change of these evolving features was different.
Contrary to the cricetids, the murids appear more conservative, as far as molar
patterns are concerned. The brachyodont-bunodont basic molar pattern prevails in
Figure 4 ( continued)
48, Mimomys (Cseria) gracilis (Ivanovce A); 49, Baranomys longidens (Gundersheim 4);
50, Ruscinomys europaeus (Orrios 3); 51, Trilophomys pyrenaicus (Serrat d'en Vacquer);
. SZ, Blancomys meini (Sarrion); 53, Cricetidae nov. gen. canterranensis (Ivanovce A);
54, Rhagapodemus frequens (Csarnota Z); 55, Kowalskia intermedia (Ivanovce A); 56,
Cricetus sp. (Ivanovce B); 57, Allocricetus sp. Clvanovce B); 58, Stephanomys minor
(Sarrion); 59, Prospalax priscus (Ivanovce A); 60, Paraethomys aff. jaegeri (Villalba
Alta); 61, Apodemus dominans (Csarn6ta Z).
6Z-75: Significant taxa of the Villanyian (Borsodia-Villanyia superzone): 6Z, 73,
Mimomys (Cseria) tornensis (Kolinany 1); 63,69, Mimomys (Mimomys) pliocaenicus
(Koliiiany 1); 64, Mimomys (Mimomvs) hajnackensis (Hajnacka 1); 65,68, Mimomys
(Mimomys) polonicus (R~bielice Kr6Iewskie); 66, 71, Villanyia exilis (Plelivec); 67, 7Z,
Borsodia petenyi (Beremend 4); 70, Mimomys (Cseria) reidi (Plesivec); 74, Cricetus
nanus (Vcelare 3); 75, Allocricetus sp. (Vcelare 4 A/7).
--76-86: Significant taxa of the Biharian (Microtus-Mimomys superzone): 76,8Z,
Pliomys lenki (Koneprusy C 718); 77,83, Mimomys (Mimomys) savini (prezletice); 78,
Allophaiomys pliocaenicus (Chlum 6); 79, Lagurodon arankae (V(:elare 4 A/7); 80,
Lemmus aff. lemmus (Koneprusy C 718); 81, Pliomys episcopalis (Koneprusy C 718);
84, Cricetus (ajor,. (Koneprusy C 718); 85, Allocricetus bursae (Koneprusy C 718); 86,
Apodemus sp. Koneprusy C 718).
87-96: Significant taxa of the Toringian (Arvicola-Microtus superzone): 87,
Arvicola terrestris; 88, Clethrionomys glareolus; 89, Microtus ratticeps; 90, Microtus
subterraneus; 91, Microtus nivalis; 9Z, Microtus gregalis; 93, Dicrostonyx torguatus;
94, Microtus arvalis-agrestis; 95, Mus musculus; 96, Rattus rattus. (The Toringian
taxa 87-94 are from the last Wiirmian glaciation deposits of the cave Dzerava skala;
taxa 95,96 are recent.)
110
Prom imomys Proml momys M i momys M i momys M i momys M l morny s M im omys M i momys .Arv lc olo A rvleo l o
i nsu lihrus rnol<l ov ieus dovok o s i occ: i tonus hoj nocktons •s pol on icu s pllocotnicus c ontlono t•rr•str ito
l 111 I
i I I I
i
M1crotus- like ti"'Imttbon d d if fer entiat i on
Rootless l!llo'er orowlnO motors
ln]~ _ -~~ e• of roots
Ouooppeoronc e of E'll'Omet tS it!'tt;.
Pres:enc.&~" af crown cem.-nt
Increase of h'I'DSO<Ionl
lncr•ose of I!O.IZJt
Mimomvs- l1ke enomelbond ditfercmtl oi iOh
Prtstnee of roots
lncr•o5inq undu la tion of the •no met crown bose
Absence of c rown cemtont
Promimomys- tike enometbond difhtrvn!l,.ojion
Promi momys Mi momys Arv • co l o

Zone l r.suti f orus occitonus ~~~~~~~nsis ~\~c0o~~~us ~~ :~~~~~~ ~~lvini J :~~~~o0n1: t•rr•st rls
Supll!:r -
zone Tr i lophomys- Rusc•nomys: Bo ..- sod i o- V l l tony•o Microtus·MitnOmys l A. r v i co t o ·M i crotus
Mom mol
Ru:s.c•nion VILtony i on 8 • ho r 1 an I Tor i n g i on
09t
Tim~ in Mo 5,0 4 ~0 3,0 1.8 0,5 0,1
..... Fig. 5 • Dental morphological changes within the Promimomys-Mimomys-Arvicola lineage recognized in the Pliocene
.....
and Quaternary of Europe. The diagrammatic representation is based on the left first lower molar shown in
occlusal (above) and buccal (below) view. Note the increasing undulation of the enamel crown base (heavy
line) and the relatively late appearance of rootless ever-growing molars.
murids. The dentition shows occasional tendencies toward an increase in crown height
and width correlated with limited morphological changes such as cusp hypsodonty
(e.g., Rhagapodemus, Occitanomys, and Ruscinomys) and stephanodonty (e.g.,
Stephanomys). With the exception of the highly specialized Microtia from Monte
Gargano, Italy (Freudenthal, 1976) with high crown "vole-like" molars, murids have not
realized any arvicolid tendencies in their molar pattern. Rootless molars are not
known in murids.
Finally, the spalacids represent a further distinct molar pattern which was
developed in connection with their fossorial habit: the so-called cylindrodonty. There
is a slightly increasing hypsodonty in the history of spalacids.
CONCLUSIONS
Biochronologic units and their limitations are the subject of conventions. A

main aspect of their use is how practical they can be applied. Therefore, the bound-
aries between biochronologic units should be defined as clearly as possible. This
precondition is realized by the biochronologic framework based on muroid rodents.
Within the succession of muroid rodents ranging from Vallesian land mammal age up to
the present, several significant faunal breaks can be recognized. It appears reasonable
to use these turning points in faunal history for the biochronologic subdivision. It is
noteworthy that two of the most important faunal changes in the history of muroid
rodents in Europe do not coincide with the boundaries of mammal ages. These faunal
breaks are situated in the late Turolian on the one hand and in the late Ruscinian on
the other hand. This observation suggests we should reconsider the number and the
limitations of mammal ages now widely used in Europe.
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115
Mein, P., Moissenet, E., and Adrover, R., 1983. L'extension et l'age des formations
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116
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117
BIOZONES OR MAMMAL UNITS? METHODS AND LIMrrS IN BIOCHRONOLOGY
Claude Guerm
Centre de Paleontolo_.gie stratigraphique et
paleoecologie associe au CNRS (UA 11)
Universite Claude Bernard-Lyon I
Z7-43 Boulevard du 11 novembre 1918
696ZZ Villeurbanne, France
INTRODUCTION
The first attempt to construct a biozonation scheme for European mammal-

bearing deposits was made by Thaler (1964). Neogene and lower Pliocene sites· of
southern France and of Spain were concerned, being situated in a series of rodent
range zones each designated after the name of its most typical site. Some years later
Heintz (1970) proposed a zonation based on Cervids for the Villafranchian of France
and Spain; that biozonation was generalized with the whole Mammal class, excepting
the Rodents, by Heintz et al. in 1974. The names given to the zones were those of the
most important concerned fossiliferous places. Besides, some studies were carried out
concerning the zonation of eastern Europe, notably by Fejfar (1976) and Fejfar and
Heinrich (1981), utilizing some rodent families.
In 1975, Mein proposed a biozonation for the whole continental Neogene of

western Europe and of some adjacent areas. Using all the available mammal families
and constantly improved since, that work is to be listed among the most important
results recently obtained in Cenozoic biostratigraphy (Mein, 1975, 1979, 1981).
From 1980 onwards, I proposed a similar zonation for the west European Pleisto-
cene, based on the same principles (Guerin, 1980, 198Z): geologically speaking, there
is no real distinction between Neogene and Quaternary. It was announced in Mein
(1979) and in Meon et al. (1979). These Pleistocene mammalian zones are named after
numbers which are numbered segmentally after the Neogene zones, the serial number
of each of Mein's zones being preceded by the MN abbreviation (for Mammals
Neogene). I suggested to call each Neogene and Pleistocene zones with MNQ letters
(for Mammals Neogene-Quaternary) preceding its number. Without difficulty- and
provisionally for macromammals only - the western European Mammal zonation was
extended to the Quaternary of the western part of the USSR (Guerin et al., 1983), then
to the Quaternary of the Middle East (Faure and Guerin, 1987; Guerin and Faure,
1988).
Although largely used, our MNQ zonation was a target to some criticism. A
number of critics aimed at putting other biozonations in place of ours. Much more
important were criticisms summoning the fundamental principles of biostratigraphy
and biozonation in the continental realm, leading those critics to replace it with a
totally different system inspired by the North American land mammal ages concept;
local faunas, regrouped in mammal units, themselves being brought together in a
succession of mammalian ages.

Before discussing these criticisms, let us remind ourselves what our mammalian
zonation is, and then examine the alleged advantages of other proposed biozonations.
THE REAL NATURE OF THE MNQ ZONES
MNQ zones are considered sometimes as chronozones, sometimes as biozones,

within that last case, a very restricted interpretation of the Oppelian zone. Surpris-
ingly enough, such a confusion between chronozone and biozone concepts is not rare.
MNQ zones are in fact another concept, totally different and following the
works of the Jurassic period stratigraphers. Why Jurassic period? Becuase not only
the creation of biostratigraphy was done within the Jurassic by Oppel, but also the
practical and theoretical work made subsequently that is appropriate to the Jurassic
period is indeed much more important than all the combined work on other periods. A
large amount of theoretical work was published in 1971 as the result of the 1967
"Colloque du Jurassique ;;_ Luxembourg," with contributions from leading scholars such
as Callomon and Donovan, Holder, Torrens, and Ziegler. When conceiving our Neogene
and Pleistocene zones, and afterwards when improving their definitions, we made
extensive use of ;their works, together with basic principles developed in Hedberg's
guide (1976).
Each MNQ zone has a triple definition:
presence of characteristic stages of evolutionary lineages; hence they are

lineage zones, or phylozones;
presence of typical mammalian associations (genera and species); hence they

·are association zones, or coenozones;
appearance of new taxa (genera and species); MNQ zones are range zones
too.
MNQ zones are, therefore, biozones pertaining simultaneously to three different

orders of biostratigraphical units. Moreover, each MNQ zone corresponds to a chrono-
zone, based on its duration. Accordingly, MNQ zones shall be considered as standard
zones as defined by Callomon and Donovan (1971): a biostratigraphic assemblage zone
limited by isochronous surfaces.
Such a definition involves the following remarks:
a standard zone is a piece of rock and not a period of time, but in terms of
field evidence the definition of the units and their recognition must be
distinguished (Callomon and Donovan, 1971);
even for the Jurassic period, few sections indeed are suitable for selection as
type section for standard zones, and standard zones are frequently, at least
in early times, defined without going as far as proposing type sections
(Torrens, 1971);
for some scholars, standard zones are at the same time subdivisions of
stages, referring to their own type profiles, and abstract formalization not
concerned with "concrete zones which depend on the range of guide fossil or
its accompanied faunas" (Holder, 1971);
because modern chronology depends on biostratigraphical methods, defini-

tions of standard zone type localities are required in that standard zones are
subjective interpretations (correlations) of objective data (Ziegler, 1971).
We, therefore, consider that MNQ zones are standard zones, are members of a
standard scale, and follow the requirements of contiguity and of commmon boundaries;
120
units of any rank do not overlap; there are no gaps between them; boundaries, which
are isochronous, between units of any one rank coincide with boundaries between units
of the next lower rank.
Consequently, MNQ zones are perfectly in accordance with the definition of

Callomon and Donovan (1971). Following the same authors, standard units must be
given a field definition to attain stability and objectivity "either in terms of appear-
ance in a stated area of the guide fossils on which recognition of the stratigraphical
unit is based, or in a described type-section by the base of the bed in which the char-
acteristic guide-fossils make their lowest appearance" (Callomon and Donovan, 1971,
P• 79).
All the preceding considerations were made for Jurassic marine sediments. By
and large these are valid for Neogene and Pleistocene continental deposits also.
Nevertheless, in my opinion, there are two restrictions.
Because we are in the continental realm, our whole standard scale shall be
restricted to the four lowest ranks, i.e., the Subzone (the smallest), the Zone,
the Mammalian Substage, and the Mammalian Stage. Mammalian age is the
time unit corresponding to a Mammal Stage standard unit. Owing to the
international procedures, real stages as biostratigraphical units of upper rank
must be used for marine sediments only.
A mammalian zone cannot be named after a fossil taxon, owing to the

complexity of its definition.
Of course, the main rules given by Callomon and Donovan remain applicable:
no obligation to use in each case the full hierarchy down to the lowest rank,
the unit not used being the lowest;
necessity for the standard units and especially the zones to be as widely
recognizable geographically as possible;
eventual use for reasons of (1) faunal distribution (paleogeographical sub-

provinces), (Z) facies (paleoecologically peculiar fossiliferous sites), (3) con-
venience (macromammals gathered directly from outcrops, micromammals
obtained by sieving), of parallel scales, defined in accordance with all the
requirements of a standard scale, and considered as regional or secondary
standard.
Figure 1 recalls, as an example, the definition of the MNQ zones for the
Pliocene and the Pleistocene of western Europe.
OTHER BIOZONATIONS AND THEIR ALLEGED INTEREST
MNQ biozonation was severely criticized by some scholars in the aim of substi-
tuting a biozonation of their own for it. This was done by Aguilar (198Z) for the
Miocene of southwestern Europe, by Cordy (198Z) for the middle and late Pleistocene
of western Europe, by Agustl et al. (1987) for what they call European early Pleisto-
cene, but is really part of the middle Pleistocene of southeastern Spain.
Incidentally, Bonifay (1984) criticized me for modeling the Pleistocene zones on

the Alpine glacial chronology, and for considering that there are in Europe, for
example, one Wurmian glacial fauna and one Mindel-Riss interglacial fauna even
though there are several geographically differentiated ones.
Figure Z shows a comparison of the MNQ zonation and zonations of Aguilar,

Cordy, and Agustl et al., together with Quaternary glacial chronology of western
Europe.
121
- 26 lti.orotue grep.li• UClioUII CU•ll• + ~ + Jliorotua oeoODOaua Pboclopua &\IUI'Orua
M:i.orot. Mlei plllli Dioroa~ + W,. + Coelo4ol'lta 'lbaluaarctoe aari tia-.
&l,uu. oab&ll• prmaiou. ftulcifer + Oriboe + -..u.tllu. pria:i.cuiu. +
aa- oab&llu. pl.lio\11 1Jrew apelMUI
x.- + ..._,. oabrer..
DQ 25 M:i.oroiua oeooaa.u. (pria:U1n)

-..<!l,. priaipai,. Mloroiua U'W'&liA + PliOIV'• leald. ul tS..U. t.pu. el.ll'cpaeua f Mlor~ aill.uiw 1
_<h,. priaiplli,. (priaiUn)
(priaUin) Coelocloa.ta •tiquiiatia aatiquitatiA + l::lorotw raUioeJ~eJ •· UTaliat
-<h..
Qod.e- a,pc-1-. ••• nartoollS.J
priaipai ..
IIIQ 24 JlloroW. breooieub orpaoeut. Llcul'ua + Aped.- + DioroaiOQP + t . - Arrtoola aapidu. 1 llaraoia aar.oi&J
lll.croiua Mlei DO&Ul.,..i• DioerorhiD\11 + Coelo4.cmia + Brt..aacaua darid.it Soriculu. 1
lliorotua srepJ.ta aartel.-ia IIIMutbu. :lDtel'MCI.iu. Unu. •paluuaJ Alopez lacopu.J
PU_,.. luaki relict\11 BaDciter taraaduaJ n..a .2!• d.....1
4lloorioetu. bUNu oorreHDIIi• Alou al.cea J Cen\la elaph\llf
nr.u. •cbersi f ~thua Saisa tatario&f Capra ibeZJ
interMCU.u.1 Coelod.onia Coelod.oa.t.a
aa.UquitaUa praac~or
Equu. ollballw pintea.ul
Bam. WrDUU
IIIIQ 23 Croouia apel . .a ilater.d.ia Plio.,. epiaoopal.ia + lll.orotw breooi._ia VW.pea TUJ.pee Y\llpuf . ,. . . prUoat
CUOD pri•oUIJ •le•
iboral.i Mtpoero. + 0.. olaoioa.iana llllpoero. pganku. f Bubalw ..-n1111ia f
•pel ..U.J Jla.uua ataillb.ei.MMia 'l'ropa.ihert• + Dioarorbi.Du. ..roJd. + Sua aorota priacu.
akinhei•uia 1 Dioarorbinu. beai toeohua
Eual.t.ooel'OII Melt krr~~Mw
IIIIQ 22 lliorotua breooian.ia med.i terra Crtoetu. aajor + Arrloola ouaUua Qqotol.,.u. olaioul.uaf Oul.o gulo 1
med.i terraneua 1 Arrtoola J&ll8otel1a + •mtrapa bon&li nr.w ot. pruarota.J lb'dzullti.llwJ
oaUan&J Pl io-v- obalind 1 Pr..ortbO&I + Uraua d.u.iJIBeri Dio .. rorhiD\18 bat t•eohw1
lliorotw acr••ti• j&aaODif DioerorhiDw etruacu. braolal"oephalw + Jla. .u:thw '\rocont.hel'11 1
Vulpn Y\llpea JaD&Oilif ~U••• lloaUrocuo boJJ&li
Oria amaoA u:Uqua SiUW aa.baobeuia + Cuai.8 etn.a.ou.
111fQ 21 Pit~ &rYalid.eu 1
P. greploid.e&f Cenua acoronatua1
Draua d.eni.Dc-ri auaaaenborueui.a J
. . . . _ • Pi~J lioroilJIJ '"plio • M.
a&rtnif IJ;yatri% + Pli~ epi•oepal.ia +
Allooricetua lnlnMJ --.bw .-rid.iOD&l.ia +
-wa
Paatbera pardu.1 P. eptlM&f
80&l'baoll.euiaJ
Dua olaotoaima
Urau. atehl1n.1 PalMolODiloa aattqw. 1 Paob,roroo1&'\a pU"rierl
+ C.~ aoOl'OD&•u. •
liiQ 20 Allophaiap pliooaeniou. ~ • J11.aooora oart1111 .Ulol'hoi._ CUelluaJ Qlia S).i&J t.pua Ua:i.d.U&I
nuu. .taJ A. p. piiJ1Void.Uf + Oob.Gtoaa + lfDolapa braob.Jpaib:ua 1 Qulo acbloaaer11 C:roou.ta a~lu&J
llioroiva ulei bwgo.ad.iMf &aw. auuaenborunab + ..._,bua &o.oiheri~a latideD&J B1ppopo•-
Soe:rgelia elbabeihUJ Cel"YW •rid.tonaliat Cenua elaphoidea + Biaont . .JOrJ Sua acrotat Capreoluaf
elapboidMJ i:uolad.ooeroa liE.• te.ruJ.ienaia/euo\enoid.ea + .llcu oarnuto~t J.. laiif'loom 1
Siuu. aieno.nia (te:rain&l at-.) Boa 1 P-liabTMu brn1.roat:r1a + DioerorhiAua Prauepoeroa nrtioomi.a f DiOft'Orlai.Jiw
.arcld.. etruaoua bracb.To•Jibl.•J D• ..roJd.
DQ 19 .lllophato.- pliocMilioua DioerorhiD.\111 etruao\111 etruaoua + Cania etr\llloua

plioo•ntcuat IUolad.ooeroa Uiraoero.t Pliob3•aa brn1.roaU1a
C.:rYUe peroleui•l C.:rYUe ~roleneia + Cuda etl'\IIIOU&J
Euolad.ooeroa tetrao~ Sua atro..u + Caaia talooa.eri
JCJ1Q 18 &luva •"t;ezaoDJ.a •enesellllie J Oallopral + cerwa phillai philiai AllOpbaiQIV'& plioouniOU8f Libr&lOMI
Croisetoceroa ra.oaua a1Dor 1 AoinotO'% + Jlagal.oria laUtrou ProouptooerM J Jlap.lorta I
Cernaa pbiliai philieit Pliotr~ + Cuaia •••••a..ta Pare4oliob.opi tlleow t CaDi a ..,.• .,..1•
Euolad.ooeroa aeneauaaia Libr&loea pl.l1ou. + sua at.:roaaii
.........,..t.
IOIQ 11 ~ plioc ..DiouaJ Bluua Guella borbomoa + .....,thwa ~~~~ridionalia Saiurua1 .Eqw. brtlaaanuat ~ibw
•iel~Gt-1• rtr•iil Leptoboa ak~topoa + £n&ow .aridicmal.iat Eualad.ooeroeJ Lep,o'bM
Cl"oizeioceroa raaoava med.iua 1 z.zaziaaiod.oruaaia + &(u.ua breaaarwa et.:ruao\I&J O&llocozoal •nesbJ.aiil
- pbilioi Yalli-ioJ llale• tb.oralit ViretailUI'Ua ach&ubi
aaolad.oo•roa ••nesemia rireii
J11Q 16 Jl:i..Jiolqlt poloniova 1 X. atel:ll.ill1 f »>UU. + 111ppar101lf OrJotol.,.. J t.pua 1 DicerorhiJNa

Dol.... h _ i..... ~pod. Tapirua + D6oerorh1Au. jeazm.reu, etru.OWJ Blu.u&J ~'ibU. gro.GYUJ
tnquaaa 1 Diovorhinua fJplophocloa + Plioirapa U'deu. ArYvDooeroa 1 Croisetooeroe 1 t.pioboai 1
je811'riratiJ .li'YarDMe~ U'd.eit hcJITcroout&J Ha.oiheri•
Crobetooeroa r..aau. raaoa\18 J
c.rona. ouaam.J Cer'fw ,arrierit
c.rn. pvd.iD. . . t.
IIIQ 15 lli.aloiQ'a oooit&DYJ Crioetw Azumoua + J;rc\ereui. . d.ODDesUlif 'froconihertual C.eri&J Alloorioetua1
ucuattd.•ut Ruaoin~· Eur7bou + DiovorhiD.ua •svbillw J Bippol.,.u.t PliopeAt&la,paJ
e\ll'Opuua 1 Trilopb.ocqa P1"D&i- B;rMDa + Eipparicu cru••l Doliohopi thecva f 'lberailurua 1 BurJ'bou 1
ou. J lowalakia iDtel'll8d.i&J Alepb.ia + Sui J Cania aiohauiJ Poataohisotheriu.J
Pliope~~ialapa d.iet.:ricbiJ Sua laiDorJ ~aolopb.od.oa. boraont.
R1ppar1on oru••l B. tbaurMf
Dioe:rorbiD.ua •sarbinw at. lit
D• .tsuelorwatoniit Cerna
wreD&to.,. f llephia l.7ri%J
Caaia ad.oDa& J lfTotereu.tea
d.ODDeauli J Draw ruaoi.DeMia
Prom_,.,• Par&boa + Dtoerar~ •subi.Dua 1

~·~··
IIIIQ 14 iD.aul.iterua
crtoetua barrierit IDw&lald.a PropotUIOOhoerua + .lllaDoua J Rhqapcd.e-J JJ.Jotereu.t.ea 1
...,... K. polonio&J Agrio'iber1• + Oen'ua aua tralia t tlr&U.J Jcrioiheri• i.Dai&D&J
Bb...apod.•- hautiaacMUh I Sua + Plioblraz' oooid.entalia .. lea pDD&Yauzi.J J'eU.a
Par &boa oord.ieri 1 .IDaaoue chriat.o11t Pliobyraz
U'VerDeuia ( priai Un) t oocidentaliaJ 'l'apirua ll.l'ftl"DeD&ia
DioerorhiD.ua •garb1Dua et. I
Fig. 1. Definition of the MNQ zones for the Plio-Pleistocene of western Europe.
122
... - -
CozdT Jgusti at .aaau- Clboial ohronoloctu
~ 1982 al. 1987 1!)82 .ll.JD- ••liluropeaD
lata Pleiatooane 26 VIII Veiohsel

1---- -
25 VII
=--
VI- IQ4
~
.... -
lilllllien
--
r---
lliddle PleiatooeDB 24 Saala
~""--'- t-
Y'
1-
-
23 llalateiD
~""--'-
22 1'1
lliDlel BJ.ater
~""--'-
21 tit-- X.Q3 Cromerien
~""--'-
20 II- aiias
I ...Q2
early Pleistocene 19
8Jid Pliocene
1-
18 ~ llaaaa
1- D17
17
1-
16 Bl.llar
I---
15
·I---
14
late lliocene 13 B 1- B 3
I---
12 D4
I---
11 Dl
I---
10 D2
I---
9 D1
lliddle l!iocene 8 C5
1-
1
I--- 13-C4
6
I---
5
C1-C2
earq lll.ocene 4
B
3
A5
Fig. z. Comparison of the MNQ zonation with concurrent biozonation and

Quaternary glacial chronologies for western Europe.
But what are these alternative zonations and their supposed advantages?
Aguilar's Zcmation
That zonation is based on Rodent associations and appearances, and on a tenta-

tive correlation with some marine deposits; endemism (at least presumed) and faunal
provincialism are taken into account, together with some supposed diachronism in the
arrivals of migrant genera and species.
123
Unfortunately Aguilar has not followed the evolution of the MNQ concept since
1975; marine correlations were already considered by Mein and myself; diachronism of
migrations is difficult to prove; the interest of endemic forms for correlation work
seems to us very limited. Moreover, a zonation restricted to the Rodentia evidently
seems to present far less possibilities than one founded on the whole Mammal class.
Lastly, Aguilar's zonation is geographically limited to southern France and to Spain.
Cordy's Zonation
Utilizing largely my faunal lists (many were published in my 1980 work and
nowhere else), Cordy's zonation is founded exactly on the same principles as for the
MNQ zonation; i.e., a combination of phylozones, coenozones, and range zones. It
leads to a strangely similar result, see Cordy (1982, table 2). That zonation was
proposed to remove some so-called defects of ours, the most important being that its
Pleistocene part would be based on the Alpine glacial chronology. Now, I explicitly
wrote in 1980 that use of any glacial chronology must be discontinued owing to
diachrony of the glaciations.
Evidentally embarrassed by the convergence of his results with mine, and

unaware of the differences between biozones, chronozones, climatozones, and
standard zones, Cordy tried to extricate himself from the difficulty by tracing
obliquely the limits of my zones, and horizontally the limits of his own!
In a word, Cordy's zonation brings us, in the best case, only a few facts whose
sole interest is to give a more complete definition of some of the middle and late
Pleistocene MNQ zones.
The Zonation of Agusti et al.
This zonation is based on the faunas of a number of Spanish sites, some of them
recently discovered in south eastern Spain.
Following a curiously reinterpreted MNQ 17 zone, four successive Rodent partial

range zones (Mimomys ostramosensis, Allophaiomys pliocaenicus, Allophaiomys
nutiensis, Arvicola cantiana) are defined. That succession is used on a frame in which
some "macromammals events" are assembled: a number of appearances and extinc-
tions of taxa (mostly genera, sometimes species) plus a rather surprising succession of
Canis etruscus, Canis etruscus etruscus, and Canis etruscus mosbachensis ,(Ag~tf et
al., 1987). Large criticisms of the MNQ zonation, already found in Moya Sola and
Menendez (1986), come with that piece of work, whose results are summarized in the
figure 1, p. 293 of Agustf et al. (1987).
For the moment the zonation of Agustf et al. (1987) needs more paleontological
work, together with a more elaborate bibliographical research; many important works
do not seem to be well known to our authors, and citations of others are sometimes
inexact (some faunal lists utilized without taking into account of the original "cf." or
"aff.," eliminating hence the uncertainty of the real presence of some taxa) or false (I
never defined MNQ 18 zone in terms of association of Allophaiomys pliocaenicus with
Procamptoceras and Megalovis).
FUNDAMENTAL OBJECTIONS AGAINST MNQ ZONATION, AND MY ANSWERS
Fundamental objections against MNQ zonation were essentially made by

Azzaroli (1977, 1982) and Daams and Freudenthal (1981). Their criticisms can be
summed up in seven points: contiguity and boundaries of the zones, permanence and
singularity of the sites, heterogeneity and diachrony of the faunas, and denomination
of the zones.
124
1. Contiguity of the Zones
Objection: A biozone is a sheet of rock characterized by the existence in its

whole thickness of a selected fossil or selected assemblage of fossils. Generally,
mammalian fossils are sporadically scattered and tend to be preserved in fossiliferous
layers or pockets separated by non-fossiliferous sheets. Therefore, mammalian bio-
zones are against the rule of contiguity.
Answer: It is well known in stratigraphy that fossils (from all kind, and not
especially mammals) usually constitute only a minor, disseminated and fractional part
of a rock stratum, and that there are barren spaces and intervals in all fossiliferous
sequences (Hedberg, 1976, p. 46). The standard biostratigraphic unit concept implies
that, for all parts of Phanerozoic time, units bounded by isochronous surfaces must
form a continuous succession without any gap or overlap. Any piece of rock should
belong to only one unit of given rank; units of the same rank do not overlap. Intervals
lacking in fossil content between two fossiliferous levels of the same standard zone
pertain to that standard zone by that very fact. There are no barren interzones in the
standard unit system (Callomon and Donovan, 1971).
Z. Boundaries of the Zones
Objection: A biozone should possess well defined boundaries. There are never
barren spaces nor sheet of time between two successive biozones, nor overlap. How-
ever, the distribution of mammalian remains is generally discontinuous, and the dis-
continuity between two successive mammalian assemblages corresponds to a much
longer duration than the time necessary for the constitution of one assemblage.
Moreover, a paleontologically different fauna can always be inserted between two
presently known faunas. Therefore, the boundaries between mammalian biozones are
fuzzy and unsteady.
Answer: That criticism, as for the preceding one, proceeds from a state of
confusion between biostratigraphical units, whose limits are generally not isochronous,
and chronostratigraphical units; the standard units system is ignored. "A standard
scale can be modified, if desired, as a result (for instance) of new information or
change of opinion, by the insertion of additional divisions in any rank of the hierarchy
without disturbing the other ranks. Only the base of such an additional division needs
to be specified" (Callomon and Donovan, 1971, p. 77). Subzones have already been
defined (MNQ 16a and b, for instance), and some others will very probably be distin-
guished in the near future (as in the case of zones MNQ 15, 17, ZO, Z6). It seems
unlikely that more drastic changes would arise in the Neogene and Quaternary conti-
nental biostratigraphy of Europe at middle or even at long range.
3. Permanence of the Fossiliferous Sites
Objection: A biozone is a sheet of rock. Now a mammalian fauna is defined

without considering the geological conditions; it could have been totally collected and
so been lost in the field; all that exists is a collection in a museum. Then a biozone
can be totally missing.
Answer: The best answer to such a criticism lies in the discerning choice of
typical sites for a standard zone; a few number of these sites will be better than a
specific one, and they must be selected among well-distributed deposits. Sites
currently chosen by us are, for example, some well located fossiliferous points of the
Montpellier and Perpignan-Roussillon fossiliferous formations for respectively MNQ 14
and MNQ 15 zones, the Vialette and Les Etouaires deposits for MNQ 16a and b sub-
zones, and the Saint-Vallier and Seneze deposits for MNQ 17 and 18 zones. They are
of adequate exposure and volume to remain permanent in the field, without any risk of
being collected in their totality and transferred to a museum. New sites exemplifying
a zone can be constantly inserted in our standard biostratigraphic scale. Stratig-
raphers of the marine deposits never choose limited nor consumable fossiliferous
outcrops as references for their standard zones, and do not seem especially confused
when discovering a new locality.
125
4. SiDgularity of the Fossiliferous Sites
Objection: Some reference localities have yielded genera and species possibly
unknown elsewhere, or very rare, or absent in most known localities of the same
presumed age. Such localities with endemic taxa, possibly giving evidence of peculiar
biotopes, would not be representative of the faunas of their age.
Answer: That objection does not take into account regular advances in paleon-
tology. New discoveries constantly show that many taxa considered as unique or rare
are open to be found in many more sites than previously believed; see, for instance,
the recent and repeated finds of Bubalus murrensis, Neogene European Hexaprotodon,
or Panthera toscana/gombaszoegensis. As for endemism, it seems to be restricted
mainly to some insular faunas. Insufficiency of present knowledge seems insufficient
reason to argue against biostratigraphy, because new discoveries are the rule, and
because the absence of a given taxon in the paleontological record is never as signifi-
cant as its presence; in that case there is no symmetry between absence and presence.
5. Heterogeneity of the Faunas
Objection: Micromammals and macromammals are both utilized for MNQ zona-
tion; that is a quality for some critics and a shortcoming for others. Well, micro- and
macromammals would be generally fossilized in sites of a different nature. Accord-
ingly, it would be necessary to constitute two different biostratigraphical scales, even
if the corresponding spans of time are partially or totally overlapping.
Answer: A first answer is that parallel scales considered as secondary or

regional standards are perfectly reasonable among standard unit system. But, more-
over, when sites adequately rich in fossil diversity are selected, reference localities
generally yield both micro- and macromammals. Alleged heterogeneity is often based
on historical and technical reasons. Micromammals were not found in many "classical"
localities because they were not looked for, and sediments were rarely sieved at the
beginning of the century. Experiences recently undertaken in some well known and
formerly excavated localities like Les Etouaires, Chagny, Vialette, have demonstrated
the presence of unexpected micromammals together with macromammals. Moreover,
faunal assemblages in MNQ reference localities shall be comprehensive in view of
accuracy, precision and ease of use.
6. Diachrony of the Faunas
Objection: Immigration and evolutionary processes would be diachronic

features. A genus or a species may appear earlier in one country than in another,
leading to the drawing of diachronic lines. A genus or a species may show in one
country a much faster evolution than elsewhere. Accordingly, it is possible to cross
the boundaries of several biozones over a limited geographic distance.
Answer: As far as we know, immigration is (geologically speaking) an instant

phenomenon, totally synchronous at a continental scale, as demonstrated by the
"Hipparion event" at the beginning of MNQ 9 zone, or by the recent rabbit - and other
feral mammals - dispersion in Australia. Flynn et al. (1984) indicate a dispersal rate
on the order of one to more than 10 km/yr, implying that large continental land
masses can be populated in some thousand years in the absence of major biogeographic
barriers. As for evolutionary processes, those leading to the emergence of new taxa
seem never to be the same in different places since the origin of one species seems to
occur in one region only. Polyphyletic theories are generally considered as inadequate
and obsolete.
7. Appellation of the Zones
Objection: Appellation of biozones with an ordinary succession of progressive

numbers would be inadequate if one or more mammal assemblage must be inserted
between two already designated faunas.
126
Answer: Such an objection is only allowable for newly worked regions, where
paleontological, biostratigraphical, and geological data are sparse and rare. That
opinion is indefensible for a region where paleontological work has been carried on for
more than one and a half century, and where the greatest change rationally expected
is no more than a subdivision of some already defined range zones, or as a slight
modification for the boundary between existing zones. Moreover, changing the
numbering of a standard zone succession if necessary does not seem especially diffi-
cult. Subdivision with numbers has the advantage of simplicity and practicality.
Another reason is that a geographical designation presents a risk if t4e unique refer-
ence locality was badly chosen. A good example is the site of Seynes, chosen not long
ago as the reference for the so-called "zone of Seynes;" the age of Seynes was first
thought to be Ruscinian but is now considered Villafranchian, resulting in a zone of
Seynes having a different age than the typical locality of Seynesl
So, from the seven points of criticism against MNQ zonation, two (contiguity and
boundaries of the zones) are refuted by the nature itself of the MNQ zones. Two
others (permanence and singularity of the fossiliferous sites) can be easily removed by
a careful choice of a small number of reference sites instead of an unique one. The
three other points (heterogeneity and diachronism of the faunas and denomination of
the zones) are highly debatable.
SOME REMARKS UPON LOCAL FAUNAS, MAMMAL UNITS, AND MAMMAL AGES
The local faunas/mammal units/mammal ages system is considered by Azzaroli

(1977, 198Z) as far better than the MNQ zonation. Such a system presents, however,
major shortcomings in each of its three ranks.
Local Faunas
A local fauna is generally no more than the fauna of one rich enough fossilif-
erous site. I do not see exactly the interest of creating a new formulation to be
utilized instead of the perfectly clear concept of fossiliferous site. But there are two
other possibilities for a local fauna: (1) to be constituted by the addition of several
scattered points of presumably the same fossiliferous deposit, with always the possi-
bility for these points to be more or less diachronous; (Z) inversely, a local fauna
should be a fossiliferous layer and not the whole faUna of a site, because many fossil-
iferous sites contain several fossiliferous layers of different ages, sometimes
separated by important barren spaces, as for instance in Stanska Skala, Mosbach,
Abbeville, Kent's Cavern. The outcome is ambiguity in designation. Moreover, the
duration of the accumulation of a fossiliferous level can be so brief that, within the
limits, arguing about so short "instants" prohibits the comparison of two different
sites; the probability for two sites to be exactly of the same age is low indeed; two
local faunas will never be contiguous in time, and boundaries between them will
always present gaps or overlaps. Lastly, the term "local" is not particularly precise
and would be better if restricted to a strict geographic sense.
Hence, the local fauna concept appears to be not especially straightforward, nor
clear, nor unambiguous. It has nothing to do with any kind of biostratigraphical unit.
Mammal Units
Following Azzaroli, a mammal unit is either an unique and sufficiently charac-

teristic local fauna, or a group of paleontologically similar local faunas. Weakness of
such a concept lies in the three following points: (1) definition is vague and the transi-
tion between the two concepts of "local fauna" and "mammal unit" is too progressive;
where is the boundary between a single unit (the local fauna) and its multiple (the
mammal unit)? (Z) variable and undefined boundaries in time and space; (3) lack of
relations between the alleged faunal list of a mammal unit and any geological, biogeo-
graphical, paleoecological, and taphonomic evidence. Moreover, naming the mammal
units after geographic names does not seem to present any advantage. Such a type of
127
determination extended to all of the world's geographical provinces and subprovinces
prevents any synthesis and tends to block the spreading of knowledge. The fact that
such geographical names are utilized in North America, South America, Australia, and
other countries is not to be taken for granted. Up to that point, why not give a name
to each unit of each continental basin, with the main result a verbal inflation, putting
a mask on what is the real problem in biostratigraphy: correlating the fossiliferous
deposits from basin to basin, then from province to province.
Hence, the mammal unit concept appears as a kind of biochron, i.e., a chrono-
stratigraphic unit not always well defined and denominated, and whose temporal
boundaries are unsteady and without contiguity.
Mammal Ages
A succession of mammal units defines a mammal age, which is largely inspired

by the North American land mammal age concept. That one was largely developed
from 1941 onwards. Defined as a characteristic assemblage of genera and refined
with such elements like first appearance of immigrant taxa and evolutionary first
occurrences, it concerns a ~on rather than a biostratigraphic Wlit, with a power of
resolution on the order of 10 years, boosted thanks to paleomagnetism to approxi-
mately 100,000 years (Woodburne, 1977; Flynn et al., 1984).
On the other hand, the highest rank in the MNQ system is the "mammalian
stage," which corresponds to a superzone. Such super zones are, for instance, Vallesian
and Turolian for the upper Miocene (MNQ 9 to 13), Ruscinian (MNQ 14 and 15) and
Villafranchian (MNQ 16 to 19) for the Pliocene and the early Pleistocene; there is no
name available for middle and late Pleistocene (MNQ ZO to Z4 and MNQ ZS and 26).
Advantages of the superzones/mammalian stages· are that they are members of a

standard biostratigraphical scale, and tied to a small number of type sections, with a
power of resolution inferior to 100,000 years, utilizing biochronological facts only. My
only reservation is with some names, like Turolian, whose distinction from Turonian is
one letter only. Such a distinction between two different names is discouraged by the
International Code of Zoological Nomenclature, which is used as a pattern for strati-
graphic nomenclature; moreover, the roughly equivalent Pikermian mammalian stage
is well defined. In the same way I disagree with the utilization of such terms as
Villanyian, Biharian, and Mosbachian, which are founded on sites with very different
layers of different ages, and are intended to concern what is no more than an unique
standard zone, i.e., MNQ 17 (middle Villafranchian) for Villanyian, MNQ 18 (upper
Villafranchian) for Biharian, MNQ ZO (base of middle Pleistocene) for Mosbachian.
However that may be, the most important unit in a standard scale is the zone.
CONCLUSION
MNQ zonation can be used for the whole European Neogene and Quaternary, and
will probably be extended in the near future to other zoogeographical provinces. It is
a system following the general rules of biostratigraphy. MNQ zones are standard
biostratigraphical units, are members of a standard scale, and follow the requirements
of contiguity and of common boundaries.
At least three other biozonations were proposed in place of ours, but none
presents any advantage.
On the other hand, seven fundamental theoretical criticisms were put forward
against .MNQ zonation in the aim to replace it with a totally different system of
mammal units, bringing together local faunas and regrouped in mammal ages. Two of
these objections (contiguity and boundaries of the zones) are refuted by the nature of
MNQ zones; two others (permanence and singularity of fossiliferous sites) can be
easily obviated; a discussion shows the weaknesses of the last three (heterogeneity and
diachrony of the faunas, denomination of the zones).
128
All things considered, MNQ zonation appears more logical and efficient than the
mammal unit system. That last one is too distant indeed from the rules of biostratig-
raphy, lacks precise boundaries in space and time, has no geological references, and
corresponds finally to a series of collections of fossils never exactly comparable and
without temporal continuity.
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Rongeurs et correlations avec l'echelle stratigraphique marine. C. R Acad. Sc.
Paris, v. Z94, p. 49-54.
Agustf, J., Moya Sqla, s., and Pons Moya, J., 1987. La sucesion de Mamiferos en el
Pleistoceneo inferior de Europa: Proposicion de una nueva escala
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(eds.), "Round Table of the W. Mediterranean Neogene." Trab. Neogeno-
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Azzaroli, A., 198Z. Remarques sur les subdivisions chronologiques du Villafranchien
Coll. intern. "Le Villafranchien mediterraJu~en." Lille, I, contributions, p. 7-14.
Bonifay, M.F., 1984. Grands Mammiferes, in Debrand-Passard, S. (ed.), "Synthese
geologique du Sud-Est de la France." Bur. rech. g€ol. min., Orleans, p. 5Z9-530.
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clature. Mem. Bur. rech. geol. min., v. 75, p. 75-81.
Cordy, J .M., 198Z. Biozonation du Quaternaire post-villafranchien continental
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Daams, R., and Freudenthal, M., 1981. Aragonian: The stage concept versus Neogene
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kontinentalen Quartars in Europa anhand von Arvicoliden (Rodentia,
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heterochrony, and temporal resolution in Cenozoic terrestrial sequences.
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nat. Hist. nat., C., XXn, 303 + Z06 p.
Heintz, E., G'!lerin, C., Martin, R., and Prat, F., 1974. Principaux gisements
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geol. min., v. 78/1, p. 169-18Z.
129
Holder, H., 1971. Grundsatzliches zur Jura-Gliederung. Mem. Bur. rech. geol. min., v.
75, P• 69-74.
Mein, P., 1975. ~esultats du groupe de travail des Vertebres. Report on activity on
the RCMNS working group (1971-1975), lUGS, regional committee on
Mediterranean Neogene stratigraphy, p. 78-81.
a
Mein, P., 1979. Rapport d'activite du groupe de travail Vertebres. Mise jour de la
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H.S./3, P• 1367-137Z.
Mein, P., 1981. Mammal zonation: introduction. Ann. geol. pays hellen., H.S./4, p.
83-88.
Meon, H., Ballesio, R., ~erin, c., and Mein, P., 1979. Approche climatologique du
Neogene superieur (Tortonien a Pleistocene moyen ancien) d'apres les faunes et
les flores d'Europe occidentale. Mem. Mus. nat. Hist. nat., B., XVU (1981), P•
18Z-195.
Moya Sqla, s., and Menendez, E., 1986. Los Artiodactilos (Bovidae y Cervidae,
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faunes et la stratigraphie du Tertiaire d'Europe. Mem. Mus. nat. Hist. nat., C.,
xvn, Z95 p.
Torrens, H.S., 1971. Standard zones of the Bathonian. Mem. Bur. rech. _geol. min., v.
7 5, p. 581-604.
Woodburne, M.O., 1977. Definition and characterizatiot) in mammalian chronostratig-
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130
LARGE MAMMAL DISPERSAL EVENTS AT THE BEGINNING
OF THE LATE vn.LAFRANCBIAN
Federico Masini and Danilo Torre
Dipartimento di Scienze della Terra

Universita di Firenze
Via la Pira 4
501Z.1, Firenze, Italy
INTRODUCTION
The Villafranchian is a biochronological unit based on a macromammal faunistic

complex. Pareto (1865) first used the term Villafranchian to refer to a fauna from
deposits cropping out near Villafranca D'Asti (Piemonte, Italy), and later extended it
to encompass the mammal faunas from the Upper and Lower Valdarno in Tuscany.
Since then the Villafranchian has been mainly used by Italian, French, and Swiss
authors to refer to many faunistic associations of western Europe.
The Villafranchian is characterized by a faunistic association made up of probos-

cideans (initially Anancus and Zygolophodon, later by Anancus and Archidiskodon, and
finally by just Archidiskodon), Leptobos, several lines of deer, some large sized
equines, and, amongst the carnivores, Pliohyaena (=Pachycrocuta), Chasmaporthetes,
Acinonyx, Megantereon, and Homotherium. The fact that of all these taxa only Equus
and Acinonyx survive today, albeit with different species, has been considered impor-
tant for the definition of the Villafranchian. Differences in the associations have
permitted the division of the Villafranchian into subunits. The various association
zones (Local Units according to some authors) have been grouped into the early,
middle, and late Villafranchian (figure 1).
Recently, some micromammal specialists have used the term Villafranchian to

refer to a unit of the rodent biochronological scale. The Villafranchian is commonly
found in the literature preceding the Villanyan. This use of the Villafranchian has
caused a certain amount of confusion. In order to insert the Villafranchian into the
micromammal biochronological scale, Fejfar and Heinrich (1981) redefined the inter-
val of the early Villafranchian with rodent associations and compressed the entire
Villafranchian into this subunit. This limitation is justifiable by the fact that Pareto
initially defined the Villafranchian in a locality that is early Villafranchian in age.
However, the Villafranchian~ Fejfar and Heinrich (1982, 1983, 1987) and Fejfar
and Horacek (1983), defined by succession Mimomys hajnackensis-polonicus, has an
upper limit whose position with respect to the end of the early Villafranchian, as
defined with macromammal associations, is unclear. In fact, if the St. Vallier
Mimomys were a M. polonicus (Chaline, 1974; Chaline and Laurin, 1984, 1986), this
limit would fall within the middle Villafranchian.
It is likewise quite probable that the limit between the Villanyan and the
Biharian does not coincide with the limit between the middle and late Villafranchian.
The Olivola Faunal Unit, which is the association characterizing the beginning of the

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:I: OLIVOLA OLIVClA
I r-- 1-----
-~--~
::;)
..... u I TEGflEN LU
LU .....
< TIGLIAN
..J
c z VILLANVAN LE CW'fT
<
c 17 ST.VALLIER ..J
2 I:
~ r <
a:
Sl.VAI.LIER
f---~
~~.-
LU
PRAETIGLIAN I .... I I I ROCANEVRA z

< 16b MONTOPOLI M<JIHPQI
I .... LU
'.~ -
> ..J I tESETOVA.IRES
..J
Q
u
..J
VILLA-
a:
r 0
..J Q
-
tJ) REUVERIAN <
..... - FRANCHI AN -
tJ)
> 16a TRIVERSA ..J
::;)
3
LA TRIVERSA c..
< VIAlHTE
~ ~~----~ I I I I lAYNA,siTE
~---~
..J
0::
RUSCINIAN RUSCINIAN 15 PERPIGNAN <
'--
PERPIGNAH
.....
Fig. 1. Pollen and mammal zonations of the middle-late
Pliocene and early Pleistocene of western Europe.
late Villafranchian in Italy (Azzaroli, 1977), includes Mimomys pliocaenicus on the

basis of the type sample collected in the Upper Valdarno (Masini and Torre, in press).
The type sample shows very advanced characteristics that render it comparable to M.
ostramosensis, an observation also confirmed by the results of the phenetic analysis
that Chaline (1984, figure 11) carried out on Mimomys. This suggests a correlation
between the Olivola Fauna and the upper part of the Villanyan. However, the
synonymy between the species M. pliocaenicus and M. ostramosensis cannot be
confirmed at present because the type sample of M. pliocaenicus is made up of a
mandible fragment with M/1 and one M/Z, which are obviously insufficient for the
statistical analyses necessary to separate closely related species of Arvicolids.
Our introduction brings up the following points:
1. It is well known that the micromammal fossils in macromammal deposits and the
macromammal fossils in micromammal deposits are often few and poorly repre-
sentative.
z. This has hindered and continues to hinder the correlation of the two biochrono-
logical successions, and makes it difficult to merge the two successions into a
single scale without a loss of information.
We, therefore, feel that it is better to keep the various scales that are based on
differing criteria separate, and to avoid using the same terms for units belonging to
different scales.
We have made these remarks to clarify our point of view regarding the meaning
of the Villafranchian before analyzing the bioevents that characterize the establish-
ment of the earliest late Villafranchian association.
We consider the association from the Olivola Unit to be typical of this period in
time. Although the Seneze fauna has also been described as a representative associa-
tion from the early late Villafranchian by some authors, we have not taken it into
132
consideration because the association known from Seneze is problematic. In fact,
there are taxa with middle Villafranchian affinities in its faunal lists, as well as taxa
comparable to forms from the late Villafranchian (e.g., Azzaroli et al., 1988). The
Seneze basin may well contain horizons dating from the middle Villafranchian through
well into the late Villafranchian. It is unfortunate that the information from the
existing fossil collections is insufficient to discriminate precisely between the various
possible horizons.
THE DISPERSAL EVENTS
Because of the dispersion both in time and space of European vertebrate

deposits, and especially Pliocene-Pleistocene deposits, individual bioevents cannot be
placed into a biochronological scale. Bioevents are generally documented in
"compressed form" in known local faunas. It is, therefore, impossible to base the
limits of the biochronological units on single events, and the units themselves are
defined on the basis of characteristic associations.
The events that distinguish the Olivola faunas from the Tegelen (Holland) and Le
Coupet (France) faunas are the arrival of Pliohyaena brevirostris, Canis etruscus,
Eucladoceros dicranios, and "Dama" nestii. Leptobos etruscus, a species character-
istic of Olivola, and Sus strozzii, which is common in the late Villafranchian, are
reported from Le Coupet (Heintz et al., 1971) in an association dominated by forms
tied to those from the St. Vallier zone. If this determination is confirmed, the
immigration of L. etruscus into western Europe is an event that clearly preceded the
others. Becauseof the problems the Seneze fauna pose, the fact that it includes the
same bovine does not confirm the information from Le Coupet. A form of Sus strozzii
with primitive characteristics, such as the conservation of the third upper incisor, is
also present at Seneze (Azzaroli et al., 1988). Though the faunas of Tegelen (older
than 1.6 Ma, on paleomagnetic data) and Le Coupet (younger than 1.9 Ma, radiometric
dating) are very similar, their relative chronological positions cannot be determined at
present. Tegelen may be more recent than Le Coupet, at least in some horizons that
contain Pantbera toscana. However, the presence of a Leptobos similar to the species
from St. Vallier conflicts with this dating. The hypothesis that _h. etruscus lived
alongside the older species for a period of time is reasonable, and the presence of very
infrequent fossils of a Leptobos comparable to the one from St. Vallier in the Seneze
fauna (Masini, unpublished data) could confirm it.
The Pliohyaena brevirostris Event
The dispersal of this large hyaenid was a widespread event that affected all of
Eurasia. Forms of this species whose morphologies vary at the subspecies level can be
found from Eurc;>pe to China in deposits dating from the early Pleistocene to the
middle Pleistocene (Howell and Petter, 1980; Qiu Zahn-Xiang, 1987). The exact time
of its arrival in the various regions bas not yet been established. Nevertheless, its
presence at Olivola and at Nihewan suggests that it spread rapidly throughout the
continent during the time when the first late Villafranchian associations were
becoming established.
Horizons dating from the early Pliocene to the Pleistocene in eastern and
southern Africa contain forms that can be referred to as Pliohyaena. Unfortunately,
it is generally impossible to establish the relations between these forms and P.
pyrenaica, P. perrieri, and E_. brevirostris. "Hyaena" bellax, collected at KromdraaiA
(type locality of the species), Sterkfontein Members 4 and 5, and at Makapan 3, is by
now generally recognized to be a large Pliohyaena that is either conspecific with P.
brevirostris or is very closely related to it phylogenetically (Howell and Petter, 1980;
Turner, 1986, 1987). The ages of Kroomdraai A, Sterkfontein, and Makapan 3 are
estimated to be about 1.5 Ma, between Z.4 and Z.8 Ma, and about 3.0 Ma (Turner,
1985). This demonstrates that the phylogenetic line of P. brevirostris was already
present in Africa from the middle Pliocene. The hypothesis that the species ~·
brevirostris originated in Africa is therefore reasonable.
133
PALEARCTIC REGION
Europe Asia Africa
~FO?
Fig. 2r---------~----------------------------------~~---.
First occurrence of Panthers "JJII'.-.Iarge hyaenids of Pliohyaena gr .. brevirostris ~

,C. s.s. (sma~l and middle si~ed true dogs of Central Asia h.iackals ""',Canis gr.falconeri
T1' . wolf hke dog - Cams etruscus b . Coyote like dog - Canis amensis ft )
:; Dispersal events discussed in the text; FO?=first occurrence datum uncertain.
'l'he Curls etruscus Event
Forms attributable to Canis are known in North America beginning in the

Hemphillian, and, surprisingly enough, from the Turolian of Spain (Canis cipio from
Concud). Other European fossils attributable to Canis have been found in the late
Ruscinian (MN 15) of southern France (Canis adoxus from St. Esteve, and C. michauxi
from Perpignan). It is doubtful that all these forms belong to the genus Canis~
stricto. The first "true" dogs appeared in North America during the Blancan. In
western Europe during the early and middle Villafranchian (MN 16 and 17) only
Nyctereutes and Vulpes were present, and there are no traces of Canis. With the
exception of the Langebaanweg find and some other reports of Canis ~ lato, the
first true dog in Africa is from deposits that are about Z.5 Ma (Turner, 1985). In
Soviet Asia, at least two forms of dogs which may have occupied a range extending to
the borders of eastern Europe have been found in deposits with fossils that correlate
134
with those from the early or middle Villafranchian of Europe. There seems to have
been a dispersal event that affected much of Eurasia during the middle Pliocene, and
affected Africa around Z.5-Z.4 Ma. Western Europe, perhaps for ecological reasons,
was excluded from this event until the times of the Olivola association, when Canis
etruscus arrived, and Seneze, where C. senezensis is present. Therefore, this dispersal
event at the beginning of the late Villafranchian seems to correspond to a final phase
of the expansion of the range of dogs in Europe.
During the succeeding faunal unit (Tasso Unit), which correlates with the early
Biharian (Torre, 1985), these early dispersal events were followed by the arrival of two
other dogs in western Europe (Torre, 1967; Azzaroli et al., 1988). Canis arnensis is
coyote-like (Kurten, 1974), while Canis falconeri is a large-sized dog. The latter is
taken to be the progenitor of Xenocyon, a characteristic form of the middle Pleisto-
cene of Eurasia. The arrival of these new forms is a large, very important dispersal
event. These dogs were distributed over a vast area, and Canis falconeri seems also to
have been present in Africa. We feel that Canis atrox from Kromdraai A and C.
africanus from Beds I and n at Olduvai (Turner, 1987; Savage, 1978) are either
conspecific with or very closely related phylogenetically to C. falconeri, and that they
arrived in Africa following this dispersal event. Ficcarelli and Torre had the opportu-
nity to examine a skull and a mandibular branch labeled Old/74-lang. K1-bone-bed 87,
collected at Olduvai, and found no significant differences between them and C.
falconeri from the Valdarno. The ages of Kromdraai A (about 1.5 Ma), and OlduVal
Bed n (about 1.4 Ma) agree with the time of arrival of C. falconeri in western Europe,
but the age of Olduvai Bed I (about 1.9 Ma) suggests an early immigration of this dog
in eastern Africa. Kurten and Crusafont (1977; fig. 1, p. 5) referred a P3/ and few
other fragmentary teeth, collected at Puebla de Valverde (Spain), to Canis cf.
falconeri, but with some hesitations. The morphology of the figured P3/ recalls the
features of some extinct felid, and we would not be surprised if it belongs to a
Dinofelis form. This find is really too doubtful to prove the occurrence of C. falconeri
in the middle Villafranchian fauna of Spain.
The Eucladoceros dicranios Event
This large deer is one of the most characteristic forms from the Olivola and
Upper Valdarno faunas. It is distinguished from Eucladoceros senezensis, which could
be its direct ancestor, by the greater complexity of the morphology of its antlers. The
systematic relations of Eucladoceros are still unclear because of the great variability
of its antlers, which are the elements that have been used the most in systematic work
(Azzaroli, 1953; Heintz, 1970). Therefore, the dispersal patterns of this species are
not very clear. In any case, it appears that E. dicranios became extinct during the
late Villafranchian, after having probably produced Eucladoceros tetraceros (Heintz,
1970). This species is no longer found in the associations from the end of the
Villafranchian, where there is instead a very large form that may belong to a distinct
lineage (De Giuli, 1987; De Giuli et al., 1987). Even though the genus Eucladoceros
was widely dispersed throughout Eurasia, the E. dicranios dispersal event was probably
limited to western Europe (Azzaroli et al., 1988).
------
The •nama• nestii Event
This medium to small sized deer probably derived from Cervus rhenanus
(=Cervus philisi), which is widespread in middle Villafranchian associations, or from
closely related forms (Azzaroli et al., 1988). Forms belonging to the "Dama" nestii
sensu lato group are well known from European late Villafranchian associations. Even
----
if these deer are well represented, there is a distinct lack of acceptable systematic
studies, and a revision is therefore in order. These forms have not been reported from
Asia. Some data from Italian localties show that "Dama" nestii sensu stricto under-
went minor modifications during the early Pleistocene (De Giuli and Masini, 1987). It
probably became extinct at the end of the Villafranchian, even if similar sized deer
that probably belonged to a very closely related lineage were still present in the
faunas of the early middle Pleistocene. The appearance of "Dama" nestii may very
well be an event limited to Europe. --- ---
135
The Leptobos etruscus Event
We feel it necessary to mention the Leptobos struscus dispersal event, even

though it precedes the ones described above, because it falls into the series of events
that lead to the fauna of the early Pleistocene.
~· etruscus was widely dispersed throughout Europe during the early Pleistocene,
especially in the southern regions. However, its presence in the associations from the
end of the Villafranchian has not been confirmed. Therefore, it probably became
extinct before the middle Pleistocene. The absence of possible ancestors to this
bovine in middle Villafranchian European deposits earlier than Le Coupet and Seneze
suggests that its origin was extra-European. With the exception of Leptobos syrticus
from Sahabi (north Africa, late Miocene-early Pliocene), which nevertheless does not
share significant characteristics with ~· etruscus, the genus Leptobos is absent from
Africa. On the other hand, the great number of species reported from numerous Asian
deposits show that Leptobos was widely dispersed and differentiated in the central-
southern area of this continent.
Some Asiatic forms, e.g., L. crassus, from the early Pleistocene of the Gonghe
Basin in China (Zheng Shaohua, 1985) and the probably related L. falconeri from the
Siwaliks, could have close phylogenetic relationships with~· etruscus. It is, therefore,
reasonable to suppose that~· etruscus was derived from Asia.
The dispersal of L. etruscus was followed by another important event that

affected the bovines: the appearance of "Leptobos" vallisami, a species belonging to
the primitive bison group, in the association of the Tasso Unit. A form that has been
referred to "L." vallisami has been recently described in association with L. crassus in
China (Zheng-Shaohua, 1985). Bison sivalensis (Upper Siwaliks) is probably also refer-
able to this group (Flerov, 1974, 1979). These data confirm the wide distribution of
these forms and show the importance of their dispersal history in the Eurasian conti-
nent.
Other dispersal events which could be tied to the early Villafranchian faunal
expansion are those of the caprids Procamptoceras and Megalovis. To date, it is not
possible to give a satisfactory definition of these events since the finds of these fossils
are rare and scant.
DISCUSSION
With the exception of the ~· etruscus event, the events we have examined here
took place between the end of the Tiglian and the beginning of the Eburonian pollen
phases. In fact, the fossils that make up the classic Tegelen Fauna were collected in
sediments that crop out in the type area of the Tiglian phase. In the Upper Valdarno,
recent studies (Bertini, in press) indicate the presence of Eburonian pollen associations
in sediments that underlie or are in part equivalent to the horizon that produced the
fossils of the Olivola zone. The date of the arrival of Canis is still uncertain, because
its presence in the Seneze Fauna raises the possibility that the event may precede the
end of the Tiglian. Paleomagnetic data gathered in the vicinity of Tegelen show that
the end of the Tiglian phase and the end of the Olduvai subchron coincide (Zagwjin,
1985). The dispersal events, therefore, seem to be consistent with the climatic
changes shown by the pollen data.
The paleomagnetic and paleoclimatic data suggest that there is a correlation

between the beginning of the late Villafranchian and the beginning of the Pleistocene
as it is defined at the Vrica section in Italy (Torre, 1987).
ACKNOWLEDGMENTS
The present work has been supported by a M.P.I. grant. The figure was drafted
by Mr. F. Landucci.
136
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138
SYNTHESIS ON THE • AQUITANIAN• LAGOMORPH AND RODENT
FAUNAS OF THE AQUITAINE BASIN (FRANCE)
Marguerite Hugueney
Centre de Paleontologie stratigraphique et

Paleoecologie de l'Universite Claude Bernard-Lyon 1
associ~ au C.N.R.S. (UA 11)
27-43 bvd du 11 novembre
F-69622 Villeurbanne Cedex, France
Michel Ringeade
Departement de Geologie
Universite de Bordeaux 1
avenue des Facultes
F-33405 Talence Cedex, France
INTRODUCTION
In the Aquitaine basin, where the stratotype of Mayer-Eymar is located, a

sequence of alternating limno-fluviatile and brackish-marine sediments documents the
Oligocene-Miocene transition. Locally, this sequence has produced mammalian faunas
(figure 1).
In the western part of the basin, there are many varied facies, representing the
changing level of the Atlantic Ocean. Mayer-Eymar (1857-58) recognizes seven units
(figure 2); to the east, it is often difficult to correlate the different beds, but it seems
that the "Marnes a Unios du Bazadais" correspond to the beginning of transgression.
Further eastward (Agenais region), the facies are less varied and can be divided as
only three units: The lacustrine beds of the "Calcaire blanc de l'Agenais" at the base,
and at the top those of the "Calcaire gris de l'Agenais;" between them are Continental
a
"Molasses" with intercalations of marine "Marnes Ostrea aginensis" (the "Marnes a
Ostrea aginensis" can also be found intercalated within the "Calcaire gris"). These
three units constitute the so-called "Trilogie agenaise."
While studying Lagomorphs and Cricetids in the mammalian faunas, Ringeade

(1978a,b) showed that the reliance on lithological correlations can lead to misinterpre-
tations. For example, the well known locality Laugnac is intercalated in the "Calcaire
gris" beds but in the section of Balizac, where the mammals are distinctly older than
those of Laugnac, the "Calcaire gris" beds are overlain by the mammal-bearing clays.
Therefore, paleontological data - and, in the continental beds, especially the
mammalian faunas - are of prime value.
Recently a calibration (using the evolutionary degree of the Globigerinoides and

Miogypsinidae-lineages) has been proposed ("grade-dates" in Gourinard et al., 1987,
and Magne et al., 1987). According to these authors, in the stratotype the base of the

Edited by E. H. Lindsay et a/.
N
+
Miocene
~ Oligocene
D Eocene
L-.-.--..1 10 k m
Fig. 1. Mammalian localities in the Aquitaine basin. 1 - Location of the

Aquitanian stratotype section; Z - Paulhiac; 3 - Balizac; 4 - Gans; 5 -
Aillas; 6- Cocumont; 7- Laugnac; 8- La Milloque; 9- La Brete; 10-
Estrepouy; 11 - Moissac 1.
Aquitanian corresponds to zz. 7 Ma and the Aquitanian-Burdigalian boundary to

Z0.6 Ma. They have also suggested an age of zz.z
± 0.4 Ma for the top of the para-
stratotype of Carry-le-Rouet (Formation biodetritique de Sausset, niveau ZZ), so that
the parastratotype almost completely antedates the Aquitanian (figure 3; Monleau et
al., 1988). The time scale of Berggren (1987) shows a difference of 1 Ma (Aquitanian =
Z3.7 Ma- Zl.8 Ma).
THE LOCALITIES AND THEm FAUNAS
Complete geographical and geological information on these localities can be

found in Feist and Ringeade (1977), Ringeade (1978b), de Bonis (1973), and Brunet
(1979). The name of the locality is followed by its number in figure 1; an asterisk (*)
following a species indicates its type locality.
La Milloque (Lot-et-Garonne) - 8: in the fluviatile beds of the Oligocene "Molasse de

l'Agenais," overlain by the "Calcaire blanc."
Lagomorphs and Rodents: Amphilagus aff. antiquus, Piezodus sp.
Theridomorpha: Archaeomys laurillardi and Issiodoromys pseudanaema, Rhizospalax,

Plesiosminthus schaubi, Adelomyarion vireti, Eucricetodon praecursor, Pseudo-
cricetodon cf. thaleri, Melissiodon quercyi, Rhodanomys sp., Pseudotheridomys
aff. parvulus, Gliravus sp., Peridyromys murinus, Pseudodryomys sp.
Other mammals (Brunet, 1979): Protapirus aginensis, Diaceratherium lamilloquense*,

Aceratherium (Mesaceratherium) aff. gaimersheimense, Anthracotherium sp.
large form, Microbunodon minimus, Palaeochoerus sp. 1 and sp. Z, Dremotherium
~ Dremotherium tolosanum, Bedenomeryx milloquensis, Hyaenodon,
"Plesictis" milloquensis, Cephalogale bonali, Pseudocyonopsis landesguei,
Haplocyon dombrowskii, Cainotherium, Cadurcotherium.
140
!::
ttl
,_
ttl
Ol Agenais
·~ STRATOTYPE (Mayer-Eymar)
"0
s... w
::>
co "Gres"
-Ca1caire 1acustre
(-20. ID 6-Fa 1un de La riey de
!::
ttl 5-Ca1caire 1acustre~
·~
Ca1caire greseux Bazas
!:: 4-Couches f1uvio-marines Q)
ttl
+-' sab1euses ou marneuses
·.- "'ttl
::> 3-Mo1asse ossifere ~ <Ji : : - !::
CT Q)
<( cons~id~s ----........,
1acustre 1 ,;;; 1· st::..., _ _ " ~~~~~~~~~~~~~~~:·~:f Ol
-21.2 2~r.!;!.es greseuses --- ~Ca1caire 4~~~~~~~~ ttl
~- 1-Argl1 es b1eues a Potami des et Cyrenes Couches sauma tres t:;J::;J~c:1~
.J1:0CJi~-~··:...:~M:t.a=:ra:n~e:a;.s&..:::....a.~:; Q)
b-1e-ues- ·~
--- Marn:es:bjan{h~es~cj"t:ine~nt~a1esr _ _ -=-_Ma_::s :J~ILLASJEicuMONTI Ol
Q) 0
!::
Q) II ~es
- ou-
vertes -
jaunes ~
u
-r'P.
I I I I 1I (I I I t l l":I~ s...
0 1-
Ol
5
1' Agena is
(continenta 1e)
r.-lL7A-;;M~IL:-7L-:: -OQ;; -U;;:;Ej
Fig. 2. Schematic representation of correlations for the different lithostratigraph ic units of the Aquitaine basin and location of
the mammal-bearin g localities; the dates in boxes represent age estimates (Ma) obtained by the method of "grade-dates"
(after Magne et al., 1987) •
.j:>.
CARRY-LE-ROUET (Aquitanian parastra&otype)
niv.
Burdigalian 23
_ _ _ _ EMERSIONFROM -22!0 -20.6 MY ( • AQUITANIAN STRATOTYPE),_ __
22------------------- -22.2 my <Maan6 &. a1.,198?)
Formation 21------------------- NN1 (Martini. 1988) • Miocene
biod6tritique 20
deSausset 19
18
17
Formation 16
bioc:lastique 1'
de Carry 1-{
13------------------- NP 2' (Martini,1988) • Oligocene
_ _ _ _ _ _ _ No{ Blow base
12
Formation 11
de Rousset 10
Formation 9
----------------- 23.7 my (Mqn6 &. a1.,198?)
parar6c:ifale 8
du Cap de Nautes ? -------------- Cap de Nautes • MP 30 (Bugueney &. al., 1988)
6
1st marine teyet '-------------Cap janet 1 • MP 30 ( Bugueney &. True:, 1976)
.
6
Formation
duRouet 3
2
Fig. 3. The Aquitanian parastratotype of Carry-le-Rouet.
142
Paulhiac (Lot-et-Garonne) - 2.: brown marls intercalated in the "Calcaire blanc de
l'Agenais" with "Helix" ramondi. Benoist (1873) thought that in the stratotype section
the "Calcaire blanc" was represented by a nodular white marl with "Helix" ramondi;
this Continental marl, overlain by the first marine bed of the Aquitanian, has never
since yielded any fossil remains; it is possible that fragments of the marine form
Natica, coming from the upper layers, were erroneously ascribed to "Helix" ramondi.
There is, therefore, no reason to correlate this Continental marl with the Calcaire
blanc.
Lagomorphs and Rodents (de Bonis, 1973): Titanomys visenoviensis, Piezodus

tomerdingensis, Plesiosminthus cf. myarion, Eucricetodon hesperius*,
Melissiodon, Rhodanomys transiens, Peridyromys murinus, Branssatoglis
concavidens, Branssatoglis fugax, Glirudinus glirulus, Heteroxerus
paulhiacensis*, Heteroxerus lavocati.
Other mammals: Dremotherium feignouxi, Amphitragulus gracilis, Oriomeryx major,

Bedenomeryx paulhiacensis*, Cephalogale ginesticus, Cephalogale ursinus,
Haplocyon elegans, Haplocyon crucians, Haplocyonopsis crassidens, Amphicyon
cf, astrei, Ysengrinia tolosana, Cynelos lemanensis, Plesictis solidus, Plesictis
palustris, Plesictis cultellatus, Plesictis sp. 1, Plesictis sp. 2., Proailurus
lemanensis, Diceratherium lemanense, Aceratherium (Mesaceratherium)
paulhiacense, Diceratherium pleuroceros, Protaceratherium cf, minutum,
Palaeotapirus cf, douvillei, Hyotherium major.
Moissac I (Tarn-et-Garonne) - 11: in the continental clays and marls separating the
"Calcaire blanc" and the "Calcaire gris" of the "Trilogie agenaise."
Rodents (de Bonis, 1973): Steneofiber, Plesiosminthus myarion, Eucricetodon cf,

?hesperius, Rhodanomys schlosseri (the five teeth were described· as Ritteneria
manca; the four lower molars show a continuous longitudinal crest with a short
mesolophid and seem to correspond better to R. schlosseri), Heteroxerus
paulhiacensis, Peridyromys cf, murinus, Peridyromys cf, brailloni, Branssatoglis
fugax, Vasseuromys priscus*, Glirudinus glirulus.
Other mammals: Cynelos rugosidens, Palaeogale minuta, Proailurus lemanensis.
GaDS (Gironde) - 4: in the "Marnes. a Unios du Bazadais" correlated with the marine
transgression of the Aquitanian.
Lagomorphs and Rodents: Titanomys visenoviensis, Steneofiber eseri, Plesiosminthus

cf, myarion, Eucricetodon gerandianus, Eucricetodon haslachensis, Pseudo-
cricetodon tbaleri, Melissiodon, Pseudotheridomys parvulus, Ritteneria molinae,
Heteroxerus paulhiacensis, Palaeosciurus feignouxi, Peridyromys murinus,
Peridyromys aff. brailloni, Armantomys sp., Glirudinus glirulus, Microdyromys
legidensis.
Other mammals: Protaceratherium minutum, Diaceratherium lemanense,

Cainotherium . laticurvatum, Amphitragulus boulangeri, A. gracilis,
Dremotherium feignouxi.
Ai1las (Gironde) - 5: in the "Marnes a Unios du Bazadais."

Lagomorphs and Rodents: Titanomys visenoviensis, Eucricetodon gerandianus,
Eucricetodon haslachensis, Pseudotheridomys parvulus, Ritteneria molinae,
Peridyromys murinus, Peridyromys aff. prosper-brailloni, Armantomys sp.
Other mammals: Protaceratherium minutum.
143
Cocumont (Lot-et-Garonne) - 6: in the "Marnes a Unios du Bazadais."
Lagomorphs and Rodents: Titanomys visenoviensis, Plesiosminthus sp., Eucricetodon
cf. gerandianus, Eucricetodon haslachensis, Pseudocricetodon cf. thaleri (slightly
larger than E.· thaleri of Coderet-3), Melissiodon sp., Pseudotheridomys parvulus,
Rhodanomys aff. oscensis, Blackia sp., Heteroxerus cf. paulhiacensis, Sciurid
indet., Myoglis cf. truyolsi, Paraglis cf. infralactorensis, Peridyromys murinus,
Peridyromys prosper-brailloni, Microdyromys legidensis.
Balizac (Gironde) - 3: clay level lying on alternating lacustrine and brackish sedi-
ments with a gray limestone at the top and overlain by the marine "Gres de Bazas"
(grade-date - 19.8 Ma). These layers show close relationship to the stratotype section
and Balizac represents a direct correlation point of marine and continental beds.
Lagomorphs and Rodents: Prolagus praevasconiensis*, Lagopsis spiracensis,

Eucricetodon cf. aquitanicus, Ritteneria manca, Pseudotheridomys parvulus,
Peridyromys occitanus, Peridyromys prosper-brailloni, Vasseuromys cf. rugosus,
Palaeosciurus feignouxi.
La Brete (Gers) - 9: in an alternating succession of lacustrine marls and limestones

overlying brackish sediments. In the vicinity of this locality, the "Calcaire gris de
l'Agenais" layers occur at a higher altitude.
Lagomorphs and Rodents: Prolagus vasconiensis, Lagopsis spiracensis, Steneofiber,

Eucricetodon cf. aquitanicus, Melissiodon cf. dominans, Pseudotheridomys
parvulus, Palaeosciurus feignouxi, Heteroxerus cf. paulhiacensis, Heteroxerus cf.
lavocati, Peridyromys occitanus, Peridyromys prosper-brailloni, Myoglis aff.
truyolsi, Branssatoglis sp., Vasseuromys cf. rugosus, Glirudinus cf. modestus,
Glirudinus bouziguensis, Microdyromys sp.
Laugnac (Lot-et-Garonne) - 7: marls intercalated in the "Calcaire gris de

l'Agenais." Tournouer (1870) mentioned Anchitherium aurelianense in the "Calcaire
gris de l'Agenais" as well as in Aillas, and yet another Anchitherium, A. radegondense,
in an upper Eocene level! These fossils, now lost, seem to have been misinterpreted
and erroneously ascribed to this genus.
Lagomorphs and Rodents (de Bonis, 1973): Amphilagus ulmensis, Prolagus

vasconiensis, Lagopsis spiracensis, (in the old faunal lists, the lagomorphs were
erroneously ascribed to Titanomys: Hiirzeler, 1945; Tobien, 1974), Steneofiber
eseri, Eucricetodon aquitanicus*, Melissiodon sp., Pseudotheridomys parvulus, no
Ritteneria (which was erroneously cited in older faunal lists), Sciurids,
Peridyromys occitanus*, Peridyromys brailloni, Vasseuromys rugosus*, Glirudinus
cf. modestus.
Other mammals: Haplocyon elegans?, ?Haplocyonoides mordax, Cynelos rugosidens,

Cynelos lemanensis, Amphicyon cf. astrei, Ysengrinia sp., Plesictis aff. solidus,
Plesictis laugnacensis, Palaeogale minuta, Plesiogale angustifrons, Amphictis
aginensis, Herpestides collectus, Proailurus lemanensis, Diaceratherium
aginense, Plesiaceratherium platyodon, Diceratherium cf. pleuroceros,
Diceratherium sp., Protaceratherium minutum, Palaeotapirus cf. douvillei,
Hyotherium major, Xenohyus venitor, Andegameryx, Dremotherium feignouxi,
Oriomeryx major, Amphitragulus gracilis, Cainotherium.
Estrepouy (Gers) - 10: in the continental "Molasse de l'Armagnac" overlying the

"Calcaire gris de l'Agenais" and the "Marnes a
Ostrea aginensis" (grade-date in the
"Marnes a Ostrea aginensis" - 19 Ma).
Lagomorphs and Rodents (Baudelot and Collier, 198Z; Bulot, 1980): Amphilagus
ulmensis, Prolagus vasconiensis, Lagopsis cadeoti, Steneofiber, Eucricetodon
infralactorensis*, Melissiodon cf. dominans, Pseudocricetodon sp., Pseudo-
theridomys parvulus, Ligerimys sp., Heteroxerus vireti, Heteroxerus sp.,
144
Palaeosciurus sp., Peridyromys occitanus, Peridyromys gregarius, ?Peridyromys
cf. aquatilis, Pseudodryomys simplicidens, Glirudinus modestus, ?Heteromyoxus
schlosseri, Paraglis infralactorensis*.
Other mammals (Ginsburg, 197 4): Peratherium sp., Plesiodimylus hurzeleri, Galerix
sp., Palaeogale minuta, Semigenetta elegans, Anchitherium aurelianense,
Rhinocerotidae indet. (small form), Rhinocerotidae indet. (large form),
Aureliachoerus aurelianensis, Cainotherium sp., Amphitragulus aurelianensis,
Lagomeryx parvulus.
BIOSTRATIGRAPHICAL RESULTS OF THE MAlN MICROMAMMAL FAMILIES
Lagomorpha
They are present in all the localities except Moissac I. They document an
important change between the lower levels where Titanomys is present and the upper
where it has disappeared and is replaced by the genera Prolagus and Lagopsis
(Ringeade, 1979). This change corresponds to the MN Za-MN Zb transition. As they
evolve very rapidly in these levels, they are most valuable to establish the succession
of the localities (figure 4).
Theridomorpha
Archaeomys and Issiodoromys exist only in La Milloque. They are characteristic

of the Oligocene levels.
!MP29! MNI i MN2a MN2b !MN3!
s: ~ ~ > n
~ s: Cl
& R Ill101 ~
c: i= I"" ;!
!i !ii
~ E§ ~ n~
~
I""
8c: ~ --1
:z: --1
101
101
--·-- ------- ------ ------- ------- ------- ------- ------- ------- ------- ~ ll'1fliqut6
------- ------- ------- ------- ------- ------- ------- -------- -·-- -·-·· ~ ubntitn6is
·--------·-------- -·----·-----·--------- --------------------- ~~
--0------------------------------------------------------------------~ap.
·------- ··-- -----· ------- ·--·--· ------- --·---- ------- ------- ------- ~ II:!IINUif~llrMiir
----------------------------------- ---------·------------------------- ~ _.....,,..._
------------------------------------------
-·-- ··--
-·----·--
·------- ------- -----· ·------ ------- -------------- -·---
-·----------- ~----
~~
------- ------ ------- ------- ------- ------- ------- ------- ------- -·-- ~ ertt*tJfl
--·--- ------ ------- -----·- ------- ------- ........ ------- ------· ------- TlNri-l:~iiMrp/111
-·-- --·-- --·-- --·---- --·- ------- --·----- --------------- ~

------- _; _____ --·-- --·--- ------- ------- ------- -·-- -·--- -·--· S1lln«11liiMr
-·-- --·--
------- ------- ------- ------- ------- ------- ------- ------- -------- ,_
--·--- ------- ------ ------- ------- ------- ------- ------- ------- ------- A~
~
------- --·-- --0-- ------- ------- ------- ------- ------- ------- -------· ~ l»>pt//iuul
-------- ____ , __ ------- -·-- --·-- --0-- ------- ------- ------- ------- ~ I/IIWfdllntl8
---------------------------- --------------- --D-- --0-- -·--- -------- ~ ~
-·-- ------- ------- -·--

-------- ------------- -·-- ------- -------------- -------
------- ------- ------ ------- ------- ------- ------- ------- ------- -·--· ~ hi-.1«/Q-..
-·--- --·--
------- --·-- ------- ------- ------· -·--· ~
~ ~
--·--- -·-- - - - --~----- - - - --·--

-·--- ----~---- - - - - -·-- --·-- -·--- ......i:.,
- ~- - - :.!::. - - ~- - - :.!:.· -.:.---~ ~_,.
.-.-.?.:_·_ -_-_-_-_-_-_-_ -_-_-_-_-_-_- ptn'fiiW
------- -·-- ------ ------- ------- ------- ------- ------- ,..,.,.,.

--0-- ------- ------- ------- ------- -------- ------- ------- ------- -------- RII«JJrJttmp liP-
------- -------
~
------- ------- --·-- ------- ------- ------- ------- ------- ------- -------- RII«JJrJttmp ~
-------- ------- ------- ------- ------- -0-- ------- ------- ------- ------- RII«JJrJttmp - -
-------- ------- ------ -·-- --·-- ------- ------- ------- -------- ------- liflfwlll* ~
------- ------- ------- ------- ------- ------- -·-- ------- ------- -------- liflfwlll* lllllfiCII
Fig. 4. Range chart of Lagomorphs and Rodents in the Aquitaine

basin (white squares - cf.).
145
Caatoridae
Moissac I, Gans, La Brete, Laugnac, Estrepouy. Always rare, they seem to be

more frequent in the upper levels.
Zapodidae
La Milloque, Paulhiac, Moissac I, Gans, Cocumont. Generally, only a few teeth

in each. locality; Plesiosminthus seems to disappear after the level of Cocumont
(MN Za-MN Zb boundary).
Cricetidae
Present in all the localities.
Adelomyarion, an Oligocene genus, is present only in La Milloque with a

primitive species of Eucricetodon, E. praecursor.
In Paulhiac and ?Moissac I (only one tooth) exists Eucricetodon hesperius; as

shown by Engesser (1985), its relations with Eucricetodon gerandianus are not clear.
The Eucricetodon gerandianus-infralactorensis lineage is very useful to establish

the succession of the localities as its size increase and morphological changes make
the distinction quite easy (figure 5). In the Bordelais localities, the succession given
by the Cricetids corresponds to that of the Lagomorpha. The few teeth of Cocumont
(only 37) are distinctly larger and more advanced than the specimens of Gans and
Aillas (Ringeade, 1978b); their size corresponds better to those of Balizac and La
Br~te but the morphology is more primitive: Transverse metalophule in all the MZ/ of
Cocumont (5 teeth); posterior metalophule (advanced character) in 98% of the MZ/ of
Balizac (5Z teeth) and in 83% in La Brete (1Z teeth).
Other lineages can be present. The Eucricetodon haslachensis lineage exists in

the lower levels (the large and primitive M/1 from Caunelles in Languedoc could be
related to this species) and, perhaps, some teeth of the Eucricetodon cetinensis
lineage as in Spain (Daams et al., 1987). The distinction between them is not always
easy.
Eomyidae
Paleontological study (all the measurements are given in millimeters; for the
nomenclature, see Alvarez-Sierra, 1987).
Rhodanomys-Ritteneria group
In the range of the Oligocene-Miocene boundary this group is well documented

and provides biostratigraphically valuable information. In the Aquitaine basin, it is
known from La Milloque, Paulhiac, Moissac I, Gans, Aillas, Cocumont, and Balizac.
All the measurements are given in millimeters.
In La Milloque the Rhodanomys group exists but is not yet studied.
Rhodanomys transiens Hugueney, 1969
Locality: Paulhiac
Material and measurements: see Aguilar, 1974
Discussion: In Paulhiac, the Rhodanomys, formerly described as R. schlosseri, is not

very different from R. transiens of Coderet-3 (Engesser, 1987) but the size distribu-
tion is slightly smaller. As the size of Rhodanomys is getting smaller with time, it
seems that Paulhiac is younger than Coderet-3.
146
M'
- simple ant.-ocone
- lransveree mehllophule )--- ( EtKTictltodt:n ~
~/wptnw
~
~~
~ cclllltJs
EtKTictltodt:n fTIItiCU'StT
- civided ant.-ocone
- poetericr mehllophule
)-- ( ~ gtlrlll1dwlus
EtKTictltodt:n IK{ll/llnicll8
( ?~~
~ i1hllltctallti.W
M2
- anteriCX" protolophule and mehllophule ( ~ gtlrlll1dwlus ,_ W:
- well developed Hnguaf ant.-ocingulum ) - - - - ( Saulcet ; Montaigu-1.-Biin* ; Gans ;
Aillaa ; Celina de Aragon
- poeteriCX" protolophule
- mCX"e Cl' 1-1ransv.-se mehllophule )---- ( ~ gtlrlll1dwlus 6. 1111.
La Chaux ; Cocumont
( - midcle size :
~ IK{ll/llnicll8
- poeteriCX" protolophule and mehllophule ( lau!Jlac*; Balizac; La Br6te; Salles ;
- ~ developed poeteriCX" ridge ~---__; Bouzigues; Schaflaueen; NavaTete
on the pwacone ( - l..ge size ; subdivision of the upper
- cisappe.-anca of Nnguaf ant.-ocingulum ( int.-naf ~ i7hlltctrr6n6is
Elllrepouy •; Univ. Ca1ollca
Fig. 5. Morphological changes in M1/ and MZ/ of Eucricetodon (* =type locality).
Rhodanomys schlosseri Deperet and Douxami, 1902
Locality: Moissac I
Material and measurements: left P4/, LS Z8 (0.86 x 0.96), fig. in de Bonis, 1973; left
M/1-Z, LS ZS (0.99 x 0.93), fig. in de Bonis, 1973; left M/1-Z, LS Z4 (0.98 x 1.02); right
M/1-Z, LS Z3 (1.00 x 0.97); right M/1, LS Z6 (0.97 x 1.10).
Description: The dimensions correspond as well to Rhodanomys schlosseri as to

Ritteneria molinae. The P4/ without mesoloph but showing a posteroloph are not rare
in R. schlosseri. In the lower molars two of the four teeth show an anterolophid with
twobranches and a short mesolophid; the longitudinal ridge is always complete and the
sinusid points obliquely backward. All these features are primitive and the few teeth
seem to correspond better to R. schlosseri than to Ritteneria.
Rhodanomys oscensis Alvarez Sierra, 1987
Locality: Cocumont
Material and measurements: (see table 1)
Description: Although overlap exists, the specimens of Cocumont are larger than
those of Gans and Aillas, especially in the length and width of M1/. The size corre-
sponds better to the Spanish form, R. oscensis of Santa Cilia (Alvarez-Sierra, 1987).
147
Table 1. Measurements of the cheek teeth of the
RhodanomGs-Rittene ria group in the
- -
Aquitaine asin.
Aillllll Length Width
min max N min max
04 0.82 1 0.83
pi 0.85 0.92 0.97 3 0.92 0.98 1.01
M1 0.90 0.92 0.97 4 0.91 1.01 1.06
M2 0.78 0.80 0.85 4 1.00 1.03 1.10
MS 0.67 0.73 0.80 2 0.82 0.86 0.90
04 1.10 1 0.71
P4 0.82 1 0.81
M1-2 0.93 0.97 1.00 8 0.95 1.01 1.10
Ms 0.75 0.80 0.83 4 0.77 0.81 0.84
G•• min
L.englh
mean max N min
Width
mean max
04 0.88 1 0.96
pi 0.88 0.90 0.95 12 0.89 0.97 1.01
M1 0.90 0.98 1.09 10 1.02 1.13 1.25
M2 0.74 0.87 0.98 20 0.96 1.09 1.21
MS 0.66 0.69 0.74 3 0.80 0.83 0.85
04 1.00 1.02 1.04 2 0.72 0.76 0.80
P4 0.80 0.84 0.89 5 0.75 0.83 0.91
M1-2 0.82 0.90 1.04 27 0.87 0.95 1.03
Ms 0.72 0.79 0.84 11 0.70 0.79 0.85
Cocumont Length Width

min miWI max N min mean max
o4 0.89 0.98 1.09 4 0.92 1.01 1.13
p4 1.00 1.06 1.15 13 1.01 1.06 1.15
M1 1.06 1.11 1.17 10 1.16 1.27 1.37
M2 0.77 0.92 1.07 14 0.98 1.10 1.28
M3 0.84 0.87 0.91 3 1.02 1.05 1.11
P4 0.86 0.92 1.03 3 0.78 0.84 0.89
M1-2 0.89 1.04 1.19 30 0.96 1.08 1.20
M3 0.82 0.89 0.95 3 0.83 0.89 0.95
P4/ 4 teeth out of 1Z have a complete longitudinal ridge with a short mesoloph but
this ridge has disappeared in 5 teeth out of lZ (33%); for this character,
Cocumont seems to be more advanced than Santa Cilia (only 19% of the P4/
without longitudinal crest).
M1-Z/ Z M1/ out of 8 show a short anteroloph, Z have only a small isolated cusp; in
the other M1/ and all the MZ/ the anteroloph is absent. Nearly SO% of the
M1/ have a short mesoloph and the longitudinal ridge is poorly developed in 7
out of 10 teeth (more than in Santa Cilia). The posteroloph is generally
absent.
In MZ/, the mesoloph is of middle length (8 teeth), short (5 teeth) or absent;

the longitudinal ridge is generally reduced (11 teeth out of 17); the posteroloph
is absent.
M3/ All the M3/ exhibit a very simple dental pattern.
P/4 As in Santa Cilia, shows a well developed longitudinal ridge; Z out of 3 have a
short posterolophid.
M/1-Z 15 teeth out of Z6 (58%) exhibit an interrupted longitudinal ridge without

mesolophid (only Z% in Santa Cilia). The posterolophid is generally absent.
M/3 Z out of 3 M/3 consist of two independent lophids.
Morphologically, the teeth of Cocumont are a little more advanced than those of
Santa Cilia (the interruption of the longitudinal ridge- advanced character- is more
148
frequent in Cocumont); it is difficult to decide if this form pertains to Rbodanomys or
to Ritteneria, since its morphology is intermediate between the two. As Cocumont
seems to be younger than Gans and Aillas (by the other means of correlation), it
demonstrates that R. oscensis must represent a form distinct from the Ritteneria
lineage.
Ritteneria molinae Alvarez Sierra, 1987
Localities: Gans, Aillas
Description of the specimens from Gans:
D4/ One tooth without anteroloph but with a long mesoloph.
P4/ They have the same morphology as in Cetina; nevertheless 4 P4/ out of 13
show a reduced mesoloph which never exists in Cetina.
M1-Z/ Generally, M1/ and MZ/ exhibit a short mesoloph and a complete longitudinal
ridge; these primitive characteristics are less frequent in Cetina.
M3/ Less than 50% of the teeth have a complete longitudinal ridge.
P/4 Corresponds well to those of Cetina.
M/1-Z The anterolophid is always present, with generally two branches; the
mesolophid is absent or reduced to a triangular cusp; the longitudinal ridge is
interrupted in 10 out of ZZ teeth (a little more than in Cetina); this material is
perhaps mainly composed of M/Z because the inferior teeth are a little smaller
than in Cetina whereas the upper molars have the same size.
M/3 In the majority of the teeth, the longitudinal ridge is interrupted.
Description of the specimens of Aillas: The few teeth agree in most features with
those of Gans.
Discussion: The size and morphology of the specimens from Gans and Aillas corre-
spond well to those of Ritteneria m.olinae from Cetina de Aragon.
In Spain, Alvarez-Sierra (1987) described Rbodanomys schlosseri in Moncalvillo and

Autol; this form is a little smaller than the older Rbodanomys transiens from Bergasa
and Paulhiac as the size of Rbodanomys is getting smaller with time. When the local-
ities are placed in chronological order, Rhodanomys oscensis from Santa Cilia and then
Ritteneria molinae have larger teeth and cannot be referred to the same lineage
without supposing a reversal in the evolutionary trends (Alvarez-Sierra, 1987, p. 39).
Hugueney (1974) referred to Rhodanomys schlosseri a small population (6 M1/, 8 MZ/,

14 M/1-Z) from Carriere Cluzel (France) characterized by a long mesoloph on all the
upper teeth except Z MZ/; the mesoloph is connected (or nearly connected) to the
paracone on 3 Ml/ and 6 MZ/ and the longitudinal ridge is interrupted before the
mesoloph so that the teeth have a typical S pattern (see Stehlin and Schaub, 1951, p.
130, fig. 189). In the M/1-Z the mesolophid is generally absent or reduced and the
longitudinal ridge is complete (only 1 tooth with an interrupted ridge and 4 with a
constriction); 1 M/3 out of 11 show an interrupted longitudinal ridge and 3 a constric-
tion. These specimens are distinctly smaller than R. schlosseri from Autol and
Moncalvillo and even smaller than Ritteneria manca from La Chaux and Alcocer 3b; in
our opinion, they represent an advanced population of the Rhodanomys transiens-
Rhodanomys schlosseri lineage characterized by a decrease in size.
149
In this hypothesis the Rhodanomys and the Ritteneria lineages are distinct even if they
share a common ancestry in Rhodanomys or another form with a less developed
mesoloph.
Some of the vertical trends are similar in the two lineages: decrease in size, loss of
the longitudinal ridges, and greater simplification. Rhodanomys seems to be char-
acterized mainly by its long mesoloph.
Is Ritteneria molinae related to Rhodanomys oscensis? As R. oscensis exists in

Cocumont, from which all the available data, stratigraphical as well as biostratigraph-
ical, are more advanced than in Gans and Aillas (where Ritteneria molinae is present),
we have to conclude that R. oscensis is not a representative of the Ritteneria lineage
but an independent contemporaneous group evolving more slowly. It is quite possible
that several lineages of eomyids appeared (or arrived) almost simultaneously at the
Oligocene-Miocene transition (as it is also the case for Glirids) in conjunction with the
great faunal change.
Ritteneria manca Stehlin and Schaub, 1951
Locality: Balizac
Material and measurements: P4/ = 0.86 x 0.89; MZ/ = 0.86 x 1.04; lower M, perhaps
M/Z = 0.8Z X 0.83.
Description: All the teeth are small and consist of two transverse and independent
ridges characteristic of R. ~· Balizac represents the last occurrence in the
Aquitaine. basin of the Rhodanomys-Ritteneria group; in the upper levels exists only
the Pseudotheridomys lineage.
Ritteneria does not exist in Laugnac (de Bonis, 1973) and was erroneously cited in
older faunal lists; but it could be because some teeth are known in Bouzigues (approx-
imately the same level as Laugnac).
Pseudotheridomys-Ligerimys group
This group is well documented in the Aquitaine basin as it ranges from La

Milloque to Estrepouy and is present in all the localities except Paulhiac and
Moissac I.
Pseudotheridomys parvulus (Schlosser, 1884)
Localities: Gans, Aillas, Cocumont, Balizac, Laugnac, Estrepouy
Material and measurements: (see .table Z)
Description: Because the samples are small and the specimens are morphologically
too similar to be distinctive, all these samples may be discussed as one. No appre-
ciable difference in size exists between them but they are a little larger than the
specimens from the stratigraphically lower locality of Saulcet. They show no signifi-
cant morphological difference but some P4/ exhibit an interrupted mesoloph in Gans (1
tooth out of 7), Cocumont (1 tooth out of 3), and Estrepouy (Z teeth out of 11, and one
without mesoloph; also one M1/ shows an interrupted mesoloph); in Balizac, Z lower
molars have a short longitudinal ridge between metalophid and mesolophid. The size
of the poor sample of Estrepouy is a little smaller than that of Laugnac and those of
the stratigraphically older localities; but it shows some advanced characteristics as a
tendency to reduction of the mesoloph in the upper teeth, and a reduction or even a
disappearing of the anterolophid in the lower molars (the labial branch of the
anterolophid is generally lost and the lingual branch encloses a little antero-internal
sinusid; the anterolophid is absent on 5 M/1-Z and Z M/3). Pseudotheridomys lacombai
Alvarez Sierra from Alcocer 3B (Spain) and La Chaux (Switzerland) - localities with
Ritteneria which has already disappeared before Estrepouy - is a small form with a
150
Table z. Measurements of the cheek teeth of the
Pseudotheridomys group in the Aquitaine
basin.
A AIIIM Lenglh Widlh
min 11111111 max N min m... max
P4 0.98 1 1.08
Ml 0.98 1.00 1.03 2 1.21 1.22 1.23
M2 0.95 1 1.25
P4 1.03 1.06 1.11 3 0.94 0.96 0.96
M1-2 1.18 1 1.07
Ms 0.92 0.92 0.93 2 0.88 0.90 0.93
a.n. Lenglh Widlh

min meen max N min 11111111 max
P4 0.87 0.94 1.03 7 0.97 1.05 1.15
M1 1.01 1.05 1.10 4 1.22 1.25 1.30
M2 0.91 0.96 1.02 3 1.16 1.18 1.21
04 0.96 1.04 1.12 2 0.66 0.74 0.83
P4 1.13 1 0.98
M1-2 1.03 1.09 1.16 6 1.02 1.08 1.11
Ms 0.98 1.00 1.03 2 0.98 0.99 1.00
Cocu_. Lenglh Wldlh
min m... max N min m... max
04 1.01 1 1.03
P4 0.95 1.00 1.04 3 1.01 1.06 1.09
M1·2 0.88 0.98 1.03 3 1.16 1.20 1.25
04 1.13 1 0.86
P4
-
1.05 1 0.88
-
M1-2 1.08 1.11 1.15 6 0.99 1.07 1.11
Blllizac Lenglh Wldlh

min max N min max
M1·2 0.92 0.96 1.01 4 1.15 1.19 1.26
P4 1.04 1.06 1.07 3 0.86 0.87 0.89
Ml-2 1.03 1.12 1.22 6 0.95 1.01 1.06
Ms 0.94 0.97 0.99 5 0.84 0.91 0.96
B La IHte Lenglh Widlh

min 11111111 max N min m..n max
04 0.99 1.06 1.15 19 0.97 1.06 1.11
P4 0.93 1.01 1.07 15 1.03 1.13 1.21
Ml 1.02 1.10 1.16 27 1.15 1.27 1.39
M2 0.87 1.01 1.15 57 1.11 1.25 uo
MS 0.78 0.83 0.86 14 0.98 1.02 1.07
04 1.13 1 0.80
P4 1.04 1.11 1.15 22 0.82 0.93 1.00
M1-2 1.03 1.15 1.26 66 0.96 1.07 1.t4
-
Ms 0.92 0.97 1.07 24 0.87 0.95 1.03
Laugnac: Lqlh Widlh

min max N min m..n max
04 1.00 1.02 1.05 2 0.96 0.99 1.01
Ml 1.07 1.09 1.12 6 1.18 1.24 1.29
M2 0.99 1.04 1.10 2 1.19 1.22 1.25
MS 0.82 1 1.00
04 1.09 1.12 1.15 5 0.69 0.74 0.81
P4 1.01 1.05 1.11 5 0.75 0.86 0.95
Ml-2 1.07 1.12 1.15 12 0.97 1.03 1.10
-
Ms 0.78 0.93 1.00 6 0.82 0.90 0.94
Eft"epour Lenglh Widlh

min max N min m..n max
P4 0.96 1.01 1.16 11 0.93 1.12 1.22
Ml 0.99 1.03 1.10 8 1.07 1.19 1.31
M2 0.92 0.98 1.05 9 1.17 1.25 1.29
MS 0.71 0.78 0.83 4" 0.84 0.94 1.00
P4 1.04 1.08 1.14 5 0.84 0.92 0.97
M1-2 0.98 1.07 1.19 14 0.88 0.99 1.08
Ms 0.92 0.95 0.99 3 0.91
151
reduction of mesoloph and anterolophid but, in these stratigraphically older levels, P.
lacombai is more advanced than the Estrepouy specimens (half of the P4/ have lost the
mesoloph); so the sample of Estrepouy seems to have no close relationship with this
species.
Pseudotheridomys aff. parvulus (Schlosser, 1884)
Locality: La Brete
Description: The upper teeth show five ridges and the sinus points obliquely forward.
D4/ The mesoloph is connected to the metacone in 11 out of 19 specimens; in only

2 teeth, it does not reach the labial border; the longitudinal ridge is complete
in 12 specimens and interrupted before the mesoloph in the others.
P4/ The mesoloph, always connected to the paracone, is interrupted in 3 teeth and
the longitudinal ridge is often interrupted before the mesoloph in unworn
teeth.
M1-2/ The mesoloph reaches the labial border and is generally connected to the
paracone except in 4 M1/ and 2 M2/. The anterior and posterior ellipse are
open labially in the unworn teeth, but the posterior tend to close more
quickly. The longitudinal ridge is interrupted before the mesoloph in 7 M1/
and 10M2/.
M3/ The first syncline is often very long and the protocone is connected to the
hypocone in all the specimens except one so that the sinus is closed. The
mesoloph is interrupted in 9 specimens.
P/4 The anterolophid meets the metalophid labially and lingually; the metalophid
is also connected to the mesolophid except in 4 specimens; the longitudinal
ridge is interrupted behind the mesolophid in 10 teeth; 2 specimens show
longitudinal ridges in the anterosinusid.
M/1-2 The anterolophid meets the metalophid generally labially and lingually; in 32
specimens it is short and restricted to the labial part of the tooth; it is more
long in 3 teeth and shows a lingual branch in 11 teeth. The mesolophid always
meets the metalophid, generally labially; the hypolophid is connected to the
posterolophid except in 13 specimens; the connection of the mesolophid to the
hypolophid by the longitudinal ridge is interrupted behind the mesolophid in 5
teeth; in most of the teeth this connection is situated near the middle line but
some specimens, especially the M/2, show a more lingual connection. The
sinusid points obliquely backward.
M/3 The closed anterosinusid is reduced and shifted lingually but it is always
present; the longitudinal ridge is interrupted in 3 teeth and the mesolophid is
incomplete in 3 teeth.
The size of the teeth is larger than in the samples from the stratigraphically upper
levels of Laugnac and Estrepouy and corresponds better to the material of
Schaffhausen (Germany), referred by Fahlbusch (1983) toR_. a££. parvulus (this locality
with Ritteneria and Eucricetodon aff. aquitanicus - distinctly more advanced than E.
gerandianus - can be roughly correlated with La Brete and Laugnac). The two popuia-
tions could be closely related and it seems possible that the large Ligerimys group
derives from such populations. In our opinion these populations have no close relation-
ship to the small and morphologically more advanced R_. lacombai (a greater number of
upper teeth without mesoloph) existing in Spain in nearly the same levels.
152
Ligerimys sp.
Locality: Estrepouy
Material and measurements: right M/Z, FSL 4691c = 1.07 x 1.19
Description: This tooth, described by Hugueney and Mein (1968) in Estrepouy led
them to ascribe the material to Ligerimys aff. lophidens. Mesolophid and hypolophid
are sub-parallel (in Pseudotheridomys, the two ridges connect on the longitudinal
ridge) so that the sinusid and the third synclinid are almost identical; the anterolophid
is reduced and shifted lingually. Morphologically, it could be compared with L.
antiguus from the German and Spanish localities of the same stratigraphic interval
(Bissingen, Navarrete del Rio, ••• ) but it range,o; among the largest teeth of these
localities and it could perhaps represent the large form L. lophidens.
Discussion: Morphologically, all these populations resemble E_. parvulus; therefore, in

the upper levels, the mesoloph is absent in some upper molars and premolars, as in
Schaffhausen (Fahlbusch, 1983). In the Aquitaine basin, Pseudotheridomys shows
variations in the population size. For example, in La Brete, the width of the upper
molars is slightly larger than in Schaffhausen, but in the more recent levels of
Laugnac and Estrepouy the teeth of Pseudotheridomys are distinctly smaller. There-
fore, we are not sure if there is only one lineage of Pseudotheridomys and the correla-
tions based on this genus seem to be hazardous, even in the same basin.
In the Languedoc region (Aguilar, 1974), the Pseudotheridomys parvulus samples of Les
Cevennes and La Paillade are as small as or even smaller than that of Saulcet
(Hugueney, 1974); on the lower molars they exhibit some remains of the secondary
ridge between metalophid and mesolophid which is typical of the primitive Pseudo-
theridomys schaubi. The Pseudotheridomys parvulus of Caunelles and Lespignan show
more advanced features (reduction of the anterosynclinid, connection of the cingula to
the vanishing cusps) but they remain small; they are smaller, especially in the width of
the teeth, than the Pseudotheridomys of the lower levels of the Aquitaine basin and
correspond better to those of Estrepouy.
Gliridae
We have to notice that the glirids of the Peridyromys prosper-brailloni group are
larger in Cocumont than in Gans and Aillas and give an additional indication for a
more recent level.
STRA'IlGRAPmCAL CONCLUSIONS AND COMPARISONS W1T11

OTHER MAMMALIAN FAUNAS
Comparison with the Aquit~Burdigalian Stratotype
Stratotype. At the base, the chronological position of Paulhiac is not clear, but
it is near the Oligocene-Miocene boundary interval. Moissac I cannot be correlated
with marine beds. Gans, Aillas, and Cocumont are not directly correlated to the
stratotype but they certainly correspond to the Aquitanian transgression. The only
direct correlation point of marine and continental beds is Balizac.
At the top, Estrepouy corresponds to the Burdigalian as it is more recent than

19 Ma (Burdigalian lower limit = Z0.6 Ma) and contains Anchitherium. Moreover,
Estrepouy seems to be a little younger than Universidad Catolica (Portugal) which has
yielded foraminifera of the N4-NS zone, corresponding to the Burdigalian.
Laugnac is overlain by layers estimated at 19 Ma but its exact age is not known;
as the presence of Anchitherium in the "Calcaire gris" is very doubtful, we have no
faunal indication to put it in the Burdigalian.
153
Parastratotype. Martini (1988) has determined an NP ZS Oligocene age for the
"Formation bioclastique de Carry-niveau 13" at nearly the same level that was
formerly correlated with the base of N4 zone by Blow. NN 1 (Miocene) is found only
in the "Formation biodetritique de .Sausset-niveau Z1," just below the level that has
yielded a grade-date of zz.z Ma. The sedimentologic study of Monleau et al. (1988)
concludes an emersion of the whole area of Marseille from ZZ to Z0.8 Ma (= the
duration of nearly the whole Aquitanian in Aquitaine). In the "Formation pararecifale
du Cap de Nautes-niveau 7" a fragmentary tooth of Plesiosminthus schaubi (Hugueney
et al., 1987) documents the Oligocene MP 30 mammalian zone (figure 3).
Comparison with the Reference Localities of the Mammalian Zcmation
La Milloque with Theridomorpha, Rhizospalax, Gliravus, Adelomyarion, and a

primitive species of Eucricetodon is distinctly older than the last Oligocene reference
locality of Coderet-couche 3 and pertains to the MP Z9 mammalian zone, reference
locality of Rickenbach (Schmidt-Kittler, 1987).
Paulhiac is the reference locality of the mammalian MN 1 zone; its fauna is

characterized by the disappearance of many typically Oligocene forms. Nevertheless,
its correlation with the Aquitanian stratotype is not clear (see above) and local
stratigraphers tend to assign it to the Oligocene. It can be placed near the Oligocene-
Miocene boundary.
Moissac I is stratigraphically above Paulhiac. Rhodanomys schlosseri and

Vasseuromys priscus, two forms that do not exist earlier, are in good accordance with
the stratigraphy and demonstrate also that this locality is older than Gans and Aillas.
Gans and Aillas with Titanomys visenoviensis, Eucricetodon gerandianus, and

Ritteneria molinae are correlated with the reference locality of Montaigu-le-Blin,
MN Za zone.
Cocumont is younger than Gans and Aillas by its stratigraphic position and the
evolutionary stage of Titanomys, Eucricetodon, and Glirids. Cocumont shows peculi-
arities among Cricetids and Eomyids (see above).
Balizac and La Brete can be referred to the reference fauna of Laugnac on the
basis of their lagomorpha (presence of the two advanced genera Prolagus and Lagopsis
with more primitive pattern than in Laugnac), Eucricetodon is also a little more
primitive than E. aquitanicus. In this level, the Ritteneria lineage becomes very rare
before disappearing in the level of Estrepouy.
Laugnac is the reference locality of the MN Zb zone; in our opinion, no decisive

fact permits, at the. moment, to correlate it with the early Burdigalian.
Estrepouy, reference locality of MN 3 zone, corresponds surely to Burdigalian.

Eucricetodon infralactorensis is distinctly larger than E. aquitanicus of Laugnac.
Ritteneria has disappeared and Ligerimys begins its evolution.
Figure 6 shows the correlations of the Aquitaine basin localities with some other
upper Oligocene and lower Miocene faunas.
ACKNOWLEDGMENTS
The authors are indebted to Dr. L. de Bonis (Paris), Dr. J. Hiirzeler and Dr. B.
Engesser (Basel) for access to collections in their care; Dr. E.H. Lindsay, Dr. L.
Ginsburg, and Dr. L.J. Flynn critically read part of the manuscript.
154
II
Fraace Sp1in Portugal Switzerland Germaay
~~ae ____ ~•npmm. _________________________ _____ ~
~ EPOUY Univ.Catolic:a
~ ------------------------------------------NIIVItl'ete--------------------------------------------------
Bouzipes Mt Vully
LAUGNAC Sdud'fhausen
LaBrtte
II!
Balizac Selles!Cb•
Lespipao Alc:oc:er 3b
-------------------------------------------------------------------------La Chaux------------------
Cauaelles Hulac:h
A ~ont
i Aillu
Limb~
MONI'AIGU Cetina
-~ --~------------------------------------------------------------------------------------------------
II
... ... Moissac: I
i_ PAULHIAC Ouzel
P)'rimont
FonwJt 11
SaulcetC6vennes Autol
Tomerclillaen
Bouday2 Roct.3,S,l
r--+---fi2n -------------------------------------------------------------------------------------weilllellburg 6
~ e. CODERET3 Broch.Fluh Roct.1,2,6,7
ill Cap Juet 1
; --i.AMiLLoouE--------------------------------------------------Ri;;b;b·---------------
0 --
Fig. 6. Correlations of the Aquitaine basin localities with other upper

Oligocene and lower Miocene localities (names of reference
localities are capitalized).
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156
THE FAUNAS AND STRATIGRAPHICAL SUBDIVISIONS OF
THE ORLEANIAN IN THE LOIRE BASIN (FRANCE)
Leonard Giusburg
Museum National d'Histoire Naturelle

Institut de Paleontologie
8 Rue de Buffon
75 Paris (5e), France
INTRODUCTION
The Orleanian is represented in the Loire Basin by the sands of the Orleanais,
~lesois, Syncline of Esvres, and the Langhian part of the Faluns of Touraine and
Anjou. I have already given a general faunal list of each particular locality in
1971-74. However, since that time, several new and important localities have been
discovered and others have been reexcavated. The study of the new faunas induced
me toward a revision of some mammal groups and new stratigraphical data appeared.
Rather than simply adding these new data, referring the latter to a heavy literature, I
prefer to give here the extensive and reevaluated faunal lists.
GEOGRAPIUCAL DISTRIBUTION, FACIES, AND LrrHOLOGY
In the Loire Basin, the Miocene begins with a lacustrine limestone called the
Beauce limestone ("Calcaire de Beauce Formation") which overlies without any discor-
dance the so-called Pithiviers, or Etampes limestone and Gatinais limestone, referred
to the Upper Oligocene. The entire series may reach 70 m in thickness and underlies
the plain of Beauce, which extends from Paris to Orleans. The Beauce limestone
occurs as far as the Blois area where intercalations of yellow or pink clay lenses found
within the town of Blois and in Selles-sur-Cher have yielded mammal faunas of
Agenian age (Ginsburg and Hugeney, 1980).
Upon this vast limestone table patches of detritic continental sands have been
deposited. They originate from the French Massif Central and are known as the Sands
of Orleanais. These are rather coarse, yellow, gray, or brown quartzofeldspathic sands
with gravel lenses. North of Orleans, this formation never exceeds 15 min thickness.
Strangely, all the richest vertebrate localities (Chilleurs-aux-Bois, Neuville-aux-

Bois, Artenay, "Aerotrain," Chevilly, Baigneaux-en-Beauce) occur in the lower first
meter of sand, just above the Beauce limestone. The composition of the faunal
assemblage alone permits to assign them a relative age. It seems thus that important
flowing waters brought from the Massif Central large quantities of sediment which
regularly eroded and reworked those fragile deposits of preceding periods and built up
new sedimentary structures as fragile as the previous ones.
South of Orleans, these sands are overlain by more argillaceous sands (Sologne
Sands) which never yielded any vertebrate remains.

Fig. 1. Fluvial and marine formations of the Orleanian of the Loire
Basin: (1) continental sands; (Z) marine Langhian Falun.
A= Artenay; Av = Avaray; B = Baigneaux-en-Beauce; Be =
Beaugency; Bo = Bossee; C = Chevilly; Ch = Chilleurs-aux-Bois;
Cht = Chitenay; D = D~nez~-sous-le-Lude; M = Manthelan; Mi =
Mirebeau; N = Neuville-aux-Bois; P = Pontigne; Pt = Pontlevoy-
Thenay; Sa = Savigne-sur-Lathan; Se = Selles-sur-Cher; Tav =
Tavers.
Farther west, similar sands occur in form of patches which do not exceed a few
meters in thickness and are deposited on the preexisting Esvres Syncline. These are
also known along the right bank of the Loire River between Beaugency and Tavers, as
well as in Pontlevoy. As occurs in the northern O:.;leans area, the mammal remains are
concentrated in lenses laying 1 to 1.5 m above the underlying Beauce limestone.
From Pontlevoy to Bauge, these coarse continental sands are overlain by a

totally different type of sediment - the "Falun." It is a consolidated sand extremely
rich in complete and fragmentary marine shells as well as large pebbles of flintstone
and Eocene, Cretaceous, and Jurassic limestone, often coated with bryozoans.
Numerous cross-bedded stratifications occur in the Falun. More than a thousand
invertebrate species have been recorded from this formation, all of Langhian age.
Two different facies have been defined in the Falun: (1) the so-called Pontilevian
(from Pontlevoy) facies, which is a shallow water, near-shore facies made of worn
shells and indicating a depth of approximately 10 m below the intertidal zone; (Z) the
so-called Savignean facies (from Savigne-sur-Lathan), a sandy limestone, very rich in
overcrusting bryozoans, which seems to have been deposited at a depth of ZO to
50 m. In the latter facies, the vertebrate remains, mainly shark teeth and mammal
remains, occur irregularly in the mass of the sediment. The thickness of the Falun
rarely exceeds 8 m.
158
THE FAUNAS
The faunal lists will be given for each locality, from the oldest to the youngest
one. This order will be substantiated below.
Lea Beilleaux
The locality of Les Beilleaux is situated in Anjou, but in the department of Indre
et Loire, parish of Savi~e-sur-Lathan. The Savigne area has long been known for its
Langhian marine deposits called "Faluns" and is the type locality of the "Savignean"
facies of the latter. In Les Beilleaux, this Falun rests upon gray continental sands
which have yielded a rich mammal fauna (Collier and Huin, 1985, 1979; Cabard et al.,
1980; Ginsburg, et al., 1981a,b). The following list has been established on the basis of
the collections mentioned in these publications.
Lagopsis cf. cadeoti Viret, 1930

Prolagus vasconiensis Viret, 1930
Amphilagus ulmensis Tobien, 1974
Melissiodon aff. dominans Dehm, 1950
Eucricetodon infralactorensis (Viret, 1930)
E. guadratum (Viret, 1930)
Ligerimys antiguus Fahlbusch, 1970
Eomys cf. rhodanicus Hugueney et Mein, 1968
Peridyromys murinus (Pomel, 1853)
Myomimus cf. aguatilis de Bruijn, 1974
Paraglis sp.
Heteroxerus sp.
Cynelos cf. helbingi (Dehm, 1950)
Cynelos schlosseri (Dehm, 1950)
Haplocyonoides mordax Hurzeler, 1940
Broiliana nobilis Dehm, 1950
Stromeriella franconica Dehm, 1950
Semigenetta elegans Dehm, 1950
Pseudaelurus transitorius Deperet, 189Z
Protaceratherium minutum (Cuvier, 18ZZ)
Diaceratherium aurelianensis (Nouel, 1866)
Aureliachoerus aurelianensis (Stehlin, 1899)
Xenohyus venitor Ginsburg, 1980
Brachyodus intermedius Mayet, 1908
Cainotherium laticurvatum ligericum Ginsburg, Huin & Locher, 1985
Amphitragulus cf. boulangeri Pomel, 1854
Lagomeryx praestans Stehlin, 1937
Procervulus dichotomus savignensis Ginsburg, Huin & Locher, 1985
Andegameryx andegaviensis Ginsburg, 1971
LaBrosse
Fossiliferous sands occur also in the La Brosse farm, at the limit between
Deneze-sous-le-Lude and Mei~e-le-Vicomte (Maine et Loire) just beneath the marine
Falun (Ginsburg and Janvier, 1970). Unfortunately, no precise section of these sands
could be made in this particular locality, since the sands lie partly below the level of
the phreatic water. When excavating this locality, we extracted the sand out of the
water and then sorted out the specimens. The fauna comprised mammal and selachian
remains, the latter being Myliobatis meridionalis, Galeocerdo aduncus, Odontaspis
cuspidata, 0. acutissima, Notidanus primigenius, that is, the most commonly
encountered species in the Faluns. Among the mammal remains, there were mainly
so-called "Ancient Burdigalian" forms. Some of them are covered with bryozoans and
have the typical shiny black patina of the specimens which have stayed in salt water
for a long time and became impregnated with manganese oxide. In contrast, others
are gray and dull in aspect, sometimes slightly brownish, like in Les Beilleaux. These
latter fossils do not seem to have been reworked. Thus, I suspect that in La Brosse
159
there may be continental sands in situ, and that only their upper part has been
disturbed or slightly reworked during the Falunian transgression, incorporating some
vertebrate remains of "Falunian" age.
All the species of the following list are represented at least by one specimen
with a mat and gray patina, regarded as characteristic of the continental deposits.
Lagopsis cadeoti Viret, 1930

Prolagus vasconiensis (Viret, 1930)
Eucricetodon infralactorensis (Viret, 1930)
Steneofiber depereti janvieri Ginsburg, 1988
Protaceratherium minutum (Cuvier, 1822)
Aureliachoerus auralianensis (Stehlin, 1899)
Xenohyus venitor Ginsburg, 1980
Brachyod.us intermedius Mayet, 1908
Cainotherium laticurvatum ligericum Ginsburg, Huin 8t Locher, 1985
Procervulus dichotomus savignensis Ginsburg, Huin 8t Locher, 1985
Lagomeryx praestans Stehlin, 1937
Oriomeryx willii Ginsburg, 1985
The Famaa of the Continental SaDds of the Esvres Syncline
The mammal faunas of Les Beileaux and LaBrosse are essentially the same, with
especially Steneofiber depereti janvieri, Xenohyus venitor, Brachyodus intermedius,
Andegameryx andegaviensis, and Lagomeryx praestans, which are unknown in younger
localities. Moreover, Les Beilleaux and La Brosse are in the same basin of Bauge-
Savig1le and in the same geological position, the continental sands being preserved
along the axis of the Esvres syncline and covered by the marine falun. Therefore, I
consider the two lists given above as representing a single fauna, that of the Miocene
continental sands of the Esvres syncline. Many of the mammal species of either
localities are reworked in the Falun, where they occur frequently. Namely,
Eucricetodon infralactorensis, Steneofiber depereti janvieri, Prolagus vasconiensis,
Lagopsis cadeoti, Amphilagus ulmensis, Cynelos cf. helbingi, Cynelos schlosseri,
Haplocyonoides. mordax, Brolliana nobilis, Stromeriella franconica, Semigenetta
elegans, Protoceratherium minutum, Diaceratherium aurelianensis, Xenohyus venitor,
Brachyodus intermedius, Lagomeryx praestans, Andegameryx andegaviensis which all
never occur above the stratigraphically lowermost levels of the Orleanian. However,
there are some species which are characteristic of these particular levels, and which
have been found in the Falun only, and not in Les Beilleaux. Thus, I consider that they
have been reworked from the same sand horizon as in Les Beilleux, and which has been
preserved in the Esvres syncline (Ginsburg et al., 198Z). These species are:
Amphicyon giganteus carnutense Antunes 8t Ginsburg, 1977

Phoberocyon hurzeleri Ginsburg, 1985
Ursavus elmensis Stehlin, 1917
Martes laevidens Dehm, 1950
Palaeogale hyaenoides Dehm, 1950
Acteocemas infans (Stehlin, 1939)
to which I add:
Plithocyon bruneti Ginsburg, 1980

Paratapirus intermedius (Filhol, 1885)
These two latter forms deserve an explanation. I described Plithocyon bruneti

(Ginsburg, 1980a) as coming from the Falun of Pontlevoy. But in the Falun of Anjou,
there are specimens of Plithocyon which reach almost the size of that of Sansan. I
determined it as Plithocyon cf. armagnacensis. The much smaller, but morphologi-
cally identical, _E. bruneti is obviously in its ascent, and I consider it as being
160
reworked. As to the specimen from Pontlevoy, it may actually come from Chitenay or
be reworked from this locality, as was the case for a molar of Brachyodus from the
Bourgeois collection, which I have pointed out earlier (Ginsburg, 1974b). Recently, E.
Cerdeno referred the tapir of the Faluns to Paratapirus intermedius (Cerdeno and
Morales, 1986). This form is mainly known in the Upper Agenian of Selles-sur-Cher
and has never been found in the Orleanais. It is thus logical to think that it has been
reworked from the oldest levels of the basin, that is the sands of the Esvres syncline,
which actually rest immediately above the Beauce limestone within which is situated
the locality of Selles-sur-Cher.
Chitenay
This locality is represented by a few and formerly collected specimens which are
distributed in the collections of Basel, Blois, Paris, and Lyon (Faculty of Sciences)
(Stehlin, 1907; Mayet, 1908). The list is as follows:
Steneofiber depereti janvieri Ginsburg, 1988

Amphicyoninae indet.
Prosantorhinus cf. germanicum (Wang, 19Z9)
Brachyodus intermedius Mayet, 1908
Cainotherium laticurvatum ligericum Ginsburg, Huin 8t Locher, 1985
Amphitragulus cf. boulangeri Pomel, 1854
Lagomeryx praestans Stehlin 1937
?Procervulus dichotomus savignensis Ginsburg, Huin 8t Locher, 1985
cf. Oriomeryx willii Ginsburg, 1985
By comparison to the list of 1974, based mainly on bibliographical data, one may
note some differences. The artiodactyls have been revised (Dineur and Ginsburg,
1986; Ginsburg et al., 1985) and I removed the Anchitherium aurelianense. The
mention of this equid was based exclusively on an upper tooth series from the Blois
area (with no other indication as to the locality). Mayet (1908) based the "blesense"
race on this specimen, which is supposed to have differed from A. aurelianense
aurelianense by a smaller size and broader and shorter premolars. Since a smaller size
usually suggests an older geological age, and since the only locality of the Blois area
which is older than the localities containing A. aurelianense aurelianense is Chitenay,
it was logical to believe that the small-sized-maxillary actually came from Chitenay.
However, Stehlin (19Z5) refuted the originality of this tooth which is only, he says, due
to the fact that it is badly reconstructed, the first molar being wrongly placed
between the two foremost preserved premolars, and thus taken for the P3/ (in modern
sense). Hence the atypical broad aspect of the premolars. As to the supposedly larger
size of the specimen, Stehlin refuted it too. I have seen the specimen and can only
confirm Stehlin's remarks and conclusions. Like him, I would place this specimen
among those from Pontlevoy.
Chilleurs-aux-Bois and Neuville-auz-Bois
The sand beds in the closely placed villages of Chilleurs and Neuville-aux-Bois
has yielded the same fauna in both localities, besides some specimens of rare carnivo-
rous forms which are represented by one specimen in each locality. Since 1974, only
Brachyodus (Dineur and Ginsburg, 1986) and Amphicyon (Antunes and Ginsburg, 1977)
have been the subject of studies. Thus, I shall give here a common list.
Steneofiber depereti Caliodorensis Ginsburg, 1971

Amphicyon giganteus carnutense Antunes and Ginsburg, 1977
Cynelos cf. helbingi (Dehm, 1950)
Ysengrinia depereti (Mayet, 1908)
Phoberocyon aurelianensis (Mayet, 1908)
161
Hemicyoninae indet.
Laphictis? vorax Dehm, 1950
Potamotherium valetoni (Geoffroy, 1833)
Pseudaelurus transitorius Deperet, 189Z
Anchitherium aurelianense (Cuvier, 18Z5)
Plesiaceratherium platyodon (Mermier, 1895)
?Prosantorhinus cf. douvillei (Osborn, 1900)
Hyotherium cf. soemmeringi Meyer, 1841
Brachyodus onoideus (Gervais, 1869)
Amphimoschus artenensis Mayet, 1908
Amphitragulus aurelianensis Mayet, 1908
Procervulus dichotomus (Gervais, 1859)
Acteocemas infans (Stehlin, 1939)
Palaeomeryx kaupi (Meyer, 1834)
Artenay
This is the richest locality of the Odeans area. The fossils formerly collected by
Nouel and Stehlin came from quarries which are located along a path linking the
Auvilliers castle to the village of Autroche (cf. Mayer, 1908, fig. 8). When I began to
excavate in 196Z, all these sandpits had disappeared. The location of the last one,
near the Auvilliers Castle could still be guessed as it was not filled and a small bush
grew in it. Around 1950, the Road and Bridge Maintenance Service, needing some sand
for building a bridge, opened a new sandpit in the same area as the old ones, along the
path between the Autroche village and Auvilliers Castle. It was also situated west to
the path and about 300 m north to the bush mentioned above in the "Stehlin quarry."
This sand being too clayish, the exploitation was given up and replaced by sand from
the Loire River. It is in this sandpit that I could carry on my own excavations until
1967. The fossils were concentrated in a north-south oriented channel, in the prolon-
gation of which is the bush of the "Stehlin quarry." Thus, we may conclude that all the
fossils from Artenay are strictly from the same layer. Since 1971, the lagomorphs
(Bucher, 198Z), carnivores (Ginsburg, 1977; Heizmann et al., 1980), and suids
(Ginsburg, 1973) have been revised and discussed.
The faunal list is as follows:
Lanthanotherium sp.
Erinaceidae sp.
Proscapanus sansaniensis (Lartet, 1851)
Trimylus neumayrianus subsequens Doben-Florin, 1964
Vespertilionidae indet.
Prolagus vasconiensis-oeningensis
Lagopsis cadeoti-penai
Amphilagus sp.
?Megacricetodon sp.
Democricetodon sp.
Melissiodon dominans Dehm, 1950
Ligerimys florancei Stehlin & Schaub, 1951
Pseudodryomys simplicidens de Bruijn, 1966
Glirudinus gracilis Dehm, 1950
Steneofiber depereti depereti Mayet, 1908
Hyainailouros sulzeri Biedermann, 1863
Amphicyon giganteus giganteus (Schinz, 18Z5)
Cynelos schlosseri (Dehm, 1950)
Ursavus brevirhinus Hofmann, 1887
Martes sainjoni Mayet, 1908
Martes muncki Roger, 1900
162
Hoplictis noueli (Mayet, 1908)
Mionictis artenensis Ginsburg, 1968
Potamotherium miocenicum (Peters, 1868)
Semigenetta? elegans Dehm, 1950
Herpestes aurelianensis Schlosser, 1888
Pseudaelurus lorteti Gaillard, 1899
Prosansanosmilus cf. peregrinus Heizmann, Ginsburg & Bulot, 1980
Gomphotherium sylvaticum Tassy, 1985
Zygolophodon turicensis (Schinz, 19Z4)
Albanohyus pigmeus (Deperet, 1892)
Brachyodus onoideus (Gervais, 1869)
Cainotherium sp.
Amphitragulus aurelianensis Mayet, 1908
Lagomeryx parvulus (Roger, 1898)
Paleomeryx kaupi Meyer, 1834
Eotragus artenensis Ginsburg & Heintz, 1968
The • Aerotram• at Chevilly
This small locality appeared during excavations for the construction of the track
for the prototype of the "Airborne Railway (Aerotrain)" (Ginsburg, 1980). At the pillar
no. 358, excavation yielded the following fauna (micromammals are studies by Bulot).

Galerix sp.
Prolagus oeningensis (Koenig, 18Z5)
Lagopsis penai Royo, 1928
Democricetodon sp.
Megacricetodon primitivus (Freudenthal, 1963)
Melissiodon aff. dominans Dehm, 1950
Pseudodryomys simplicidens de Bruijn, 1966
Miodryomys biradicus Mayer, 1979
Ligerimys florancei Stehlin & Schaub, 1951
Heteroxerus rubricati Crusafont, Villalta & Truyols, 1955
Steneofiber depereti depereti Mayet, 1908
Pseudaelurus cf. romieviensis Roman & Viret, 1934
Gomphotherium cf. angustidens (Cuvier, 1817)
Deinotherium cuvieri Kaup, 1831
Prosantorhinus cf. douvillei (Osborn, 1900)
Brachypotherium brachypus (Lartet, 1837)
Cainotherium miocenicum Crusafont, Villalta & Truyols, 1955
Dorcatherium guntianum Meyer, 1847
Dorcatherium cf. naui Kaup, 1833
Amphimoschus artensis Mayet, 1908
Lagomeryx minimus (Toula, 1884)
Eotragus cf. sansaniensis (Lartet, 1851)
Baigneauz-en-Beauce
Apparently, all fossils labeled "Baigneaux-en-Beauce" come from the same

quarry (sandpit), situated at a crossroad, southeast of the village. I excavated it from
time to time, between 1961 and 1971. The faunal list is as follows, but .the carnivores
163
(Ginsburg and Bulot, 198Z) and rhinoceroses (Ginsburg and Bulot, 1984) were revised
after the discovery of the Bezian fauna (Gers).

Steneofiber depereti carnutense Ginsburg, 1971
Cynelos helbingi (Dehm, 1950)
Pseudocyon sansaniensis Lartet, 1851
Agnotherium aff. grivense (Viret, 19Z9)
Hemicyon stehlini Hurzeler, 1944
Ursavus cf. brevirhinus Hofmann, 1887
Hoplictis noueli (Mayet, 1908)
Trocharion cf. albanense Major, 1903
Pseudaelurus romieviensis Roman & Viret, 1934
Prosansanosmilus peregrinus Heizmann, Ginsburg & Bulot, 1980
Gomphotherium angustidens (Cuvier, 1817)
Zygolophodon turiciensis (Schinz, 18Z4)
Deinotherium sp.
Plesiaceratherium lumiarense Antunes & Ginsburg, 1984
Prosantorhinus germanicus (Wang, 18Z9)
Lartetotherium sansaniensis (Lartet, 1851)
Hyotherium soemmeringi Meyer, 1834
Bunolistriodon lockharti (Pomel, 1848)
Procervulus dichotomus dichotomus (Gervias, 1859)
Palaeomeryx kaupi Meyer, 1834
The Scmds of Beaugency-Tavers
This formation extends over about ZO km along the right bank of the Loire,
between Baule and Avaray. Recently, excavations were undertaken again in Tavers
(Ginsburg et al., 1987), and thanks to the construction of the highway, a small excava-
tion could be made in Le Bardon (Ginsburg, 1974b). The fauna of this sand is as
follows:

Pseudaelurus romieviensis Roman and Viret, 1934
? Archaeobelodon filholi (Frick, 1933)
Deinotherium cuvieri Kaup 1831
Prosantorhinus douvillei (Osborn, 1900)
Gaindatherium rexmanueli Antunes & Ginsburg, 1984
Bunolistriodon lockharti (Pomel, 1848)
Dorcatherium naui Kaup, 1833
Amphimoschus pontileviensis Bourgeois, 1873
Procervulus dicbotomus (Gervais, 1859)
164
Dicrocerus elegans parviceros Ginsburg, 1967
Palaeomeryx cf. lathanensis Ginsburg, 1985
'1be Sauds of Pont1evoy-Thenay
In the area of the two neighboring villages of Pontlevoy and Thenay, all the
quarries show, over a thickness of several meters, a fine yellow "Falun" which is more
or less consolidated and extremely rich in marine, often worn shells. Below this
"Falun," there is about one meter of clayish, rather coarse, and slightly ocreous conti-
nental sands, which rest in their turn upon the Beauce limestone. These sands are
often eroded by the overlying "Falun" which, in extreme cases, rests directly on the
Beauce limestone. There the surface of the latter is perforated by numerous litho-
fagous organisms (pholads and lithodomes), indicating that the Falun sea was very
shallow in this distal part of the gulf.
The mammal remains in the so-called Pontlevoy fauna comes from either the
continental sands, or the lower part of the Falun, as clearly indicated by Mayet (op.
cit., p. Z51-Z5Z, Z54-Z60). In the captions to the text-figures and plates of his work,
Mayet has indicated, when he could do so, the level where the specimen came from
(sand or falun). But how is one to trace the origin of the other specimens? One can,
on one hand, use Mayet's remark that the ocreous patina, which is frequently observed
in the specimens from the sand, does not occur on the specimens from the Falun. The
latter are always more black. On the other hand, one notices that in the Falun of
Anjou, the mammal fauna is represented almost exclusively by isolated teeth, the tide
and waves having dissociated the teeth. In contrast, the Bourgeois collection (studied
by Mayet) comprises a large number of more or less complete mandibles coming from
the Falun, but these have often retained the ocreous patina which characterizes the
specimens from the continental sand. These specimens are, for instance, the mandible
of Trochictis zibethoides mut. florancei (Mayet, 1908, Pl. 1Z:13) mentioned as coming
from the Falun. The same applies to the mandible of Macrotherium grande (op. cit.,
Pl. 10:13), the cracks of which still retain some typical grayish sand grains.
Thus, one can admit that most of the taxa in the so-called "Pontlevoy fauna" of
the Bourgeois collection come from the continental sands which underlie the Falun,
even if some specimens have been found reworked in the base of the Falun.
The isolated teeth found in the Falun remain of doubtful origin as they may
come undistinctively from the Falun or the sand as well. The only criterion· of distinc-
tion, the patina, is valid only if the specimens are strongly ocreous. For this reason, I
shall not refer to the Pontlevoy Pliopithecus in the following list, as it was repre-
sented in the Bourgeois collection only by an isolated tooth, the patina of which
cannot be assessed since the specimen was destroyed during World Warn.
Moreover, the faunas from the Falun and the sands are the more difficult to
distinguish as no major time gap exists between the two formations. Worse, the Falun
may sometimes be interbedded in the sand and the bone-bearing gravels (Ginsburg,
1980c). In the small Falun-pit of Les Grandes Noues, 600 m WNW of the hamlet of
l'Epini~re, between Thenay and Choussy, Denizot (19Z7, p. 4Z7, fig. Z7) showed that
the bone-bearing gravels hollow out sands containing marine fossils and are overlain by
the Falun. Similarly, in the southeastern corner of the quarry of the water reservoir
north of Pontlevoy (= "Perroche quarry" in Denizot and Four a Chaux Quarry in
Macaire, 1977) one could observe, some years ago, that the surface of the Beauce
limestone, which was perforated by pholads, had been overlain by about 1 m of grayish
sands containing bones and teeth of continental mammals; the trench showed at the
base the gray sand containing bones and sandstone blocks and, above, the mass of the
Falun, which is several meters thick. It appears thus that in the beginning of the
transgression of the Falun sea, slight oscillations of the ground may have caused varia-
tions of the coastline in the deepest part of the gulf, hence these locally alternating
marine and continental sediments and wear-preservation phenomena.
165
The fauna of the sands is as follows:

Prolagus cf. oeningensis (Koenig, 18Z5)
Cynelos bohemicus (Schlosser, 1899)
Hemicyon stehlini Hurzeler, 1944
Thaumastocyon bourgeoisi Stehlin & Helbing, 19Z5
Martes muncki Roger, 1900
Hoplictis florancei (Mayer, 1908)
Semigenetta repelini Helbing, 1937
Herpestes aurelianensis (Schlosser, 1888)
Protictitherium gaillardi (Major, 1903)
Pseudaelurus romieviensis Roman & Viret, 1934
Zygolophodon turiciensis (Schinz, 18Z4)
Archaeobelodon filholi (Frick, 1933)
Deinotherium cf. bavaricum Meyer, 1831
Chalicotherium grande (Lartet, 1851)
Brachypotherium brach}lus (Lartet, 1837)
Prosantorhinus douvillei Osborn, 1900)
Bunolistridon lockharti (Pomel, 1848)
Procervulus dichotomus dichotomus (Gervais, 1859)
Stephanocemas elegantulus (Roger, 1904)
Palaeomeryx lathanensis Ginsburg, 1985
The I.anghian Fahm of Touraine and Anjou
The first transgression of the Falun sea reached the bottom of the Pontlevoy
gulf. The Faluns have yielded mammals in Pontlevoy, Thenay (Mayet, op. cit.; Stehlin,
op. cit.; Ginsburg and Sen, 1977; Sen and Makinsky, 1983), at Bossee and Manthelan
(Mayet and Lecointre, 1909) in the gulf of Loches, and at Mirabeau in the Poitiers
Gulf, as well as north of the Loire in the area of Savi~e-sur-Lathan and the Bauge
area. In these two latter contiguous areas, the Falun contains the remains of
contemporaneous mammals mixed with older ones, reworked from the underlying
continental sands. These reworked sands are mainly of the same age as the fauna of
Les Beilleaux, but some specimens from Pont Boutard, in the eastern end of the same
basin, are suggestive of the age of Artenay or slightly younger. Consequently, I
removed from the faunal list of the Faluns all of the obviously reworked mammals. I
withdrew also from the list the suid Conohyus simorrensis that we found also in the
a
upper part of the Falun which is well known as "FalUQ Area." These upper beds of
the Falun contain also Hipparion primigenium and Deinotherium giganteum and have
to be correlated to the Falun of Do!J(;-la-Fontaine which has yielded a typical
Vallesian fauna (Ginsburg et al., 1979). The emended list is as follows:
Pliopithecus piveteaui Hurzeler, 19 54

Plesiodimylus sp.
Lanthanotherium sp.
Proscapanus cf. sansaniensis (Lartet, 1851)
Galerix sp.
166
Soricidae indet.
Spermophilinus bredai (Meyer, 1848)
Miopetaurista cf. ~ (Mein, 1958)
Miodyromys cf. aegircii Baudelot & Mein, 197Z
Gliridae ind.
Monosaulax minutus (Meyer, 1838)
Cricetodon aureus Mein & Freudenthal, 1971
Megacricetodon collongensis (Mein, 1958)
Megacricetodon cf. ~ (Mein, 1958)
Democricetodon cf. mutilus Fahlbusch, 1964
Eumyarion weinfurteri (Schaub & Zapfe, 1953)
ADchitheriomys w1edemanni (Roger, 1885)
Prolagus oeningensis (Koenig, 1825)
Lagopsis penai Royo, 1928
Amphicyon giganteus giganteus (Schinz, 1825)
Agnotherium grivense (Viret, 1929)
Cynelos bohemicus (Schlosser, 1899)
Pseudocyon sansaniensis Lartet, 1851
Pseudarctos bavaricus Schlosser, 1899
Thaumastocyon bourgeoisi Stehlin & Helbing, 1925
Alopecocyon goeriachensis (Toula, 1884)
Hemicyon cf. sansaniensis Lartet, 1851
Plithocyon cf. armagnacensis Ginsburg, 1955
Martes cf. filholi (Dep~ret, 1887)
Martes cf. muncki Roger, 1900
Ischyrictis zibethoides (Blainville, 1841)
Hoplictis cf. florancei (Mayet, 1908)
Mionictis dubia (Blainville, 1841)
?Potamotherium miocenicum (Peters, 1868)
Taxodon sansaniensis Lartet, 1851
Trocharion albanense Major, 1903
Semigenetta cf. sansaniensis (Lartet, 1851)
Protictitherium gaillardi (Major, 1903)
Protictitherium crassum (Dep~ret, 1892)
Pseudaelurus transitorius De~ret, 1892
Pseudaelurus quadridentatus (Blainville, 1841)
Sansanosmilus palmidens (Blainville, 1841)
Zygolophodon turicensis (Schinz, 1825)
Deinotherium bavaricum Meyer, 1831
Anchitherium aurelianense (Cuvier, 1825)
Chalicotherium grande (Lartet, 1851)
Prosantorhinus douvillei (Osborn, 1900)
Lartetotherium sansaniensis (Lartet, 1851)
Hispanotherium matritensis (Prado, 1854)
Bunolistridon lockharti (Pomel, 1848)
Taucanamo sansaniensis (Lartet, 1951)
?Cainotherium cf. miocenicum Crusafont, Villalta & Truyols, 1955
Dorcatherium peneckei Hofmann, 1892
Procervulus dichotomus dichotomus (Gervais, 1859)
Stephanocemas elegantulus (Roger, 1904)
Lagomeryx pumilio (Roger, 1898)
167
Palaeomeryx lathanensis Ginsburg, 1985
Eotragus sansaniensis (Lartet, 1851)
BIOSTRA'TIGRAPHY
Besides the Pontlevoy area, where the Falun rests directly on the Pontlevoy
sand, and some localities in Anjou (Savigm!-sur-Lathan, Deneze-sous-le-Lude) where
the Falun rests on continental sands of Les Beilleaux type, the fossiliferous localities
are scattered, and the patches of gray sand in which they are located rest directly
upon the Beauce limestone. Consequently, the study of the faunas is the only way of
assigning relative ages to these localities. The faunas differ from one another by the
degree of evolution of the species of the same lineage, the immigrant species, and the
extinctions. The latter criterion is difficult to apply because the species disappear
progressively, not always in the same time in all places. Sometimes, however, an
abundant species may become extinct suddenly when eliminated by a dominant
immigrant. In such a case, the appearance of the immigrant is enough to characterize
the fauna. Moreover, the cases of sudden extinction are far from more frequent than
immigrations. Extinctions will not be considered in establishing the chronological
order proposed here.
The Cbitenay-Les Beilleaux Group
The faunas of these two localities are almost identical and characterized by the
presence of Brachyodus intermedius and the first antler-bearing cervids, as well as the
absence of Anchitherium. Their relative age is difficult to assess, Chitenay being
much poorer than Les Beilleaux. However, the two artiodactyla Andegameryx
andegaviensis and Lagomeryx praestans are slightly larger in Chitenay, suggesting a
younger age for the latter locality. In this case, one may admit that the immigrants in
Les Beilleaux should logically occur in Chitenay as well. By comparison with the
Table 1. Stratigraphical distributions of the Primates, Lagomorpha, and

large Rodentia in the Orleanian of the Loire Basin. Bei = Les
Beilleaux biostratigraphical; Ch = Chilleurs-aux-Bois; Art =
Artenay; Aer = Aerotrain; Bx = Baigneaux-en-Beauce; Ta = Tavers;
Ptl = Pontlevoy (Pontlevoy sands); fal = Faluns of Touraine and
Anjou.
MN3 MN4 MNS

Bel [h Art Aer ex Tav Ptl fa I
Pliopithecus piveteaui ? +
Prolagus vasc::::n~ens.:.s .,.
II
vasconiensis-oeningensis +
II
oeningensis + + cf. +
Lagopsis cadeoti
II
cadeoti-penai "'" +
+ +
penai
.,.
II
Amphilagus ulmensis cf.

Steneofiber deperet~ Janv~er~
.. caliodorensis "'" +
II II
depereti + +
II II
carnutense + + + +
Monosaulax minutus +
Anchitheriomys wiedemanni +
168
immediately older localities (Laugnac, Selles-sur-Cher), the common fauna of
Chitenay-Les Beilleaux is characterized by the following immigrants:
Brachyodus intermedius
Lagomeryx praestans
Broiliana nobilis
Stromeriella franconica
Semigenetta elegans
and the autochthonous taxa which have evolved in the same area:
Eucricetodon infralactorensis
Steneofiber depereti janvieri
Pseudotheridomys antiquus
Lagopsis cadeoti
Amphicyon giganteus carnutense
Cynelos cf, helbingi
Pseudaelurus transitorius
Oriomeryx willii
The Chilleurs-aux-Bois, Neuville-aux-Bois Groups
These localities are characterized by the following immigrants:
Anchitherium aurelianense
Amphimoschus artenensis
and new forms which arose through local evolution:
Brachyodus onoideus
Steneofiber depereti caliodorensis
Palaeomeryx kaupi
The Artenay Group
The Artenay fauna is characterized by the following immigrants:
Gomphotherium sylvaticum
Zygolophodon turicensis
Hyainailouros sulzeri
Hoplictis noueli
Mionictis artensis
Prosansanosmilus peregrinus
Albanohyus pigmeus
Eotragus artenensis
and new autochthonous forms:
Steneofiber depereti depereti

Proscapanus oasconiensis-oeningensis
Lagopsis cadeoti-penai
The • Airborne Railway• (~erotrain) Group
In this locality, several immigrants occur:
Deinotherium cuvieri
Brachypotherium brachypus
Dorcatherium guntianum
Dorcatherium cf. naui
169
Table 2. Stratigraphical distribution of the Creodonta and terrestrial Carnivora
in the Orleanian of the Loire Basin.
MN3 MN4 MNS

Bei (h Art Aer ex Tav p tl fal
Hyainailouros sulzeri + + + + +
Cynelos schlosseri + +
Cynelos cf .. helbingi +
II
helbingi ? +
II
bohemicus + +
Amphicyon major +
l!mphicyon giganteus carnutense + +
II II
giganteus
.,. + + + + +
Ysengrinia depereti
Pseudocyon sansaniensis +
Pseudarctos bavaricus +
Alopecocyon goeriachensis +
Agnotherium grivense + +
Haplocyonoides mordax +
+ +
Thaumastocyon bourgeoisi
Hemicyon stehlini
.,. 1"
II
cf.sansaniensis +
Plithocyon bruneti +
II
cf.armagnacensis +
Phoberocyon huerzeleri +
II
aurelianensis +
Ursavus elmensis +
II
brevirhinus + +
Martes laevl.dens +
Martes sainjoni +
Martes muncki + + cf.
Martes filholi cf.
Palaeogale hyaenoides + + +
Laphictis ? vorax +
Hoplictis noueli + +
II
florancei + ct.
Ischyrictis zibethoides +
Potamotherium valetoni +
+
...+ + +
II
miocenicum
Mionictl.s artenensis + ...
II
dubia +
Broiliana nobilis +
Stromeriella franconica +
Taxodon sansaniensis +
Trocharion albanense cf. +
Semigenetta elegans + cf.
II
repelini +
II
sansaniensis cf.
Herpestes aurelianensis ... ...
Protictitberium gaillardi + +
Protictitherium crassum +
Pseudaelurus transitorius + +
II
lorteti +
II
romieviensis ct. + + +
II
quadridentatus +
Prosansanosmilus peregrinus ? +
Sansanosmilus palmidens +
170
and the species belonging to autochthonous lineages:
Prolagus oeningensis
Lagopsis penai
?Pseudailurus cf. P. romieviensis
Eotragus cf. sansaniensis
The Baignea.ur-en-Beauce Group
In the Baigneaux-en-Beauce Group, one can note, by comparison to the "Airborne

Railway" Group, the occurrence of the three immigrants:
Bunolistridon lockharti
Ischyrictis zibethoides
Lagomeryx minimus
and one new autochthonous form:
Steneofiber depereti carnutense
The Beaugency-Tavers Sand Group
Here, two new immigrants appear:
Dicrocerus elegans parviceros

Gaindatherium rexmanuelli
and two autochthonous new species:
Amphimoschus pontilenviensis
Palaeomeryx cf. lathanensis
The Pontlevoy-Thenay Sand Group
Three forms occur here by immigration:
Thaumastocyon bourgeoisi
Ischyrictis zybethoides
Chalicotherium grande
and one by autochthonous lineage:
Hoplictis florancei
The Faluns of Touraine and Anjou
One may note the appearance of some immigrants:
Pseudarctos bavaricus
Taucanamo sansaniensis
Hispanotherium matritensis
and, likely,
Pliopithecus antiquus
171
Table 3. Stratigraphical distribution of the Proboscidea, Perissodactyla, and
Artiodactyla in the Orleanian of the Loire Basin.
MNJ. MN4 MNS

Bei Ch Art Aer BX Tav Ptl fa I
Gomphotherium sylvaticum +
" angustidens ? + + + +
Zyll_olophodon turicensis + + + +
Deinotherium cuvieri + ? +
" bavaricum ct. +
Anchitherium aurelianense + + + + + +
Chalicotherium grande + +
Paratapirus intermedius +
Protaceratherium minutum + +
Plesiaceratherium platyodon +
" lumiarense + + +
Diaceratherium aurelianensis + + +
Prosantorhinus gerrnanicus ? +
" douvillei d. d. + + + +
Brachypotherium brachypus + + + + +
Lartetotherium sansaniensis + +
Gaindatherium rexmanueli +
Hispanotherium matritensis +
Aureliachoerus aurelianensis + + + + + + +
Hyotherium soernrneringi d. + + + +
Xenohyus venitor +
Bunolistriodon lockharti + + + +
Taucanamo sansaniensis +
Albanohyus pigmeus + ?
Brachyodus intermedius +
" onoideus + +
Cainotherium laticurvatum ligericum +
" miocenicum +
Dorcatherium guntianum + + + +
Dorcatherium naui d. + + + +
Dorcatherium peneckei +
Arnphimoschus artenensis + + + +
" pontileviensis + + +
Arnphitragulus boulangeri cf. + +
Arnphitragulus aurelianensis + +
Procervulus dichotomus savignensis +
" " dichotomus ct. ? + + + + ct.
Dicrocerus elegans parviceros + + +
Acteocemas infans + +
Lagomeryx pumilio +
Lagomeryx parvulus + + +
Lagomeryx minimus + + + + +
Lagomeryx praestans +
Stephanocemas elegantulus + +
Oriomeryx willii +
Palaeomeryx kaupi + + + +
" lathanensis + + +
Andegameryx andegaviensis +
Eotragus artenens.1s +
" sansaniensis d. +
172
The new forms arising from autochthonous lineages are:
Hemicyon cf. sansaniensis

Mionictis dubia
Semigenetta sansaniensis
Pseudaelurus quadridentatus
BIOZONATION
I have tried to gather the localities in the following manner:
The localities of Les Beilleaux, Chitenay, and Chilleurs form a rather homoge-
neous unit with the association of Anchitherium and Brachyodus, the earliest antler-
bearing "cervids," and the absence of proboscideans. The MN3 zone has been defined
by the occurrence of Brachyodus nr. onoideus and Anchitherium. Our revision seems
to show that these two immigrations have not been exactly synchronous. Since
Anchitherium is the more universally known form being comparable in importance to
Hipparion, which characterizes the upper Miocene, should we consider that the local-
ities containing Brachyodus nr. onoideus but without Anchitherium are younger than
MN3? The ensemble of the immigrants (Broiliana, Stromeriella, Semigenetta, and
above all the first antler-bearing "cervid") suggests the contrary. Thus, we may
propose to define the base of a biozone by the first occurrence of one of the major
immigrants which have first been used to define the biozone in question. Thus, we
would have here a MN3a biozone characterized by the occurrence of Brachyodus
intermedius and Lagomeryx praestans and a MN3b biozone with Anchitherium
aurelianensis.
The younger faunas form a new group, with the first occurrence of the
proboscideans. This very important event has been called "Proboscidean datum"
(Bernor et al., 1987). The Artenay fauna represents a first step with the occurrence of
mastodonts, those of the "Airborne Railway" and Baigneaux a second step, with the
arrival of Deinotherium. The "Airborne Railway" locality marks also the arrival of
Dorcatherium. Moreover, Bunolistriodon lockharti occurs in Baigneaux. The absence
of suids in the "Airborne Railway" locality could be regarded as insignificant and due
to local conditions. However, Steneofiber from the same locality is still ~· depereti
depereti as in Artenay, and not the more advanced subspecies of Baigneaux. Conse-
quently, we propose to place Artenay in the MN4a biozone, following here Mein (1975,
1977) and Fahlbusch (1976), and contrary to Mein (1979).
The localities of the Blois area group differ from the preceding ones by the first
occurrence of the genera Dicrocerus, Chalicotherium, and Pliopithecus. However,
these rare forms have not been found in all localities. Only Dicrocerus elegans
parviceros, found in Tavers, Thenay and the Faluns is fairly characteristic of the
earliest members of this group of localities, in contrast to the faunas of the Orleans
area. The faunas of the Beaugency-Tavers group have long been undistinguishable
from those of Pontlevoy-Thenay, but recently the discovery of two immigrant
Rhinocerotidae coming from the lberic peninsula has enabled us to make a distinction
(Ginsburg et al., 1987). In the Lisbon Basin, Gaindatherium appears suddenly in the
level R3 of the classical classification of Telles Antunes (1971), and is immediately
very abundant. Then, it becomes very rare and, in the younger R4 level,
Hispanotherium appears massively and shortly. The same sequence occurs in the Loire
Basin. Gaindatherium appears suddenly in the Beaugency-Tavers locality and
disappears, then Hispanotherium occurs in the Faluns of Anjou. As it is not possible to
intercalate the Beaugency-Tavers sands between the Pontlevoy-Thenay sands and the
Falun, I shall consider them as being older. Moreover, the presence of Chalicotherium
in Thenay and not in Tavers supports this chronological order.
Finally, the separation of the faunas of Pontlevoy-Thenay from that of the

Faluns is substantiated by a stratigraphical argument, the superposition of the two
formations, and also local evolutionary changes in various lineages. The ensemble of
173
these three locality groups form the MN5 biozone, the base of which is marked by the
immigration of Dicrocerus elegans.
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175
Stehlin, H.G., 1925. Catalogue des ossements de Mammiferes tertiaires de la
collection Bourgeois a!'Ecole de Pont-Levoy (Loir-et-Che r). Bull. Soc. Hist. nat.
Loir-et-Che r, v. 18, p. 77-277.
Tassy, P., 1985. La place des mastodontes miocenes de !'Ancien Monde dans la
Phylogenie des Proboscidea (Mammalia): hypotheses et conjectures. These
Univ. Paris 6, Paris, 861 p.
176
A PRELIMINARY MAMMAL ZONATION OF THE
UPPER MARINE MOLASSE OF SWITZERLAND
Burkart Engesser
Naturhistorisches Museum
Augustinergasse 2
CH-4001 Basel, Switzerland
At the "International Symposium on Mammalian Biostratigraphy and Paleo-

ecology of the European Paleogene" in 1987 in Mainz, Nestor Mayo and I have pre-
sented a biozonation of the Lower Freshwater Molasse based on mammals (see
Engesser and Mayo, 1987). In this biozonation the Lower Freshwater Molasse was
subdivided into 18 mammal zones. In contrast to previous zonations, complete
mammal faunas were used, not only one or two families. These zones are based on the
evolutionary level of well known evolutionary lines, the first and the last record of
certain taxa, and on typical associations. Each zone is defined by a so-called
"reference fauna." In addition, as many as possible molasse faunas of about the same
age were integrated into each zone.
Subsequently, I have constructed the zonation of the Upper Marine Molasse,

which is the subject of this paper, following the same system. In comparison with the
zonation of the Lower Freshwater Molasse, which is based on more than 100 mammal
faunas, the zonation of the Upper Marine Molasse is much less consolidated. As a
matter of fact, the interval of time represented by the Upper Marine Molasse (about 3
million years) is less than half that of the Lower Freshwater Molasse.
This Upper Marine Molasse zonation is very fragmentary at this moment; how-
ever, it is nevertheless an advancement, because until a few years ago the range of
the Upper Marine Molasse was a blank spot within the mammal zonation of the Swiss
Tertiary.
The sediments of this time period in Switzerland are mainly marine; therefore
incidental findings of mammals are rare. And for a long time it was considered hope-
less to look for them. Indeed some faunas of large mammals were known as Briittelen,
Bucheggberg, La Moliere, and others (see Stehlin, 1914; Rutsch and Schl\ichter, 1973),
but other than marine mammals such as cetaceans and sirenids, these faunas contained
only rhinoceroses, tapirs, cervids, suids, and carnivores which are very hard to cali-
brate chronologically.
This situation changed only in this decade when private collectors of shark teeth
accidentally came across small mammals in their washed samples•. Subsequently, it
turned out that small mammal remains were not rare at all in these, marine sands. At
the same time we managed also to find several small mammal faunas in freshwater
sediments, which were temporal equivalents of the Upper Marine Molasse. By this
means a dozen small mammal faunas of the range of the Upper Marine Molasse came
together. These sometimes very meager faunas permit subdivision of the Upper
Marine Molasse into four mammal zones.

Edited by E,R Lindsay et aL
Table 1. The preliminary mammal zonation of the Upper Marine Molasse. (The
assemblage zone of Mt. Vully 1 belongs to the Low.er Freshwater Molasse
that of Hirschthal to the Upper Freshwater Molasse.) '
Reference faunas Characteristic Correlations

of the Assemblage First Record Last Record Typical Associations within the
Taxa
Zones Molasse
Llgerimys florancel Anomalomys

OBERKULM
HIRSCHTHAL Megacricetodon Democricetodon ligerimys Ligerimys +Anomalomys
cf.b011rgeoisl EBIKON (Hallen)
Prolagus cf.oenlngensis
Megscrlcetodon ROGGLISWIL
Megacricetodon
WATTWIL ligsrlmys nov.sp. Euniyarion Melissiodon Eumylirlon +MeUsslodon ? STAFFELBACH
Gomphotherium ? WIKON
HINTERSTEINBRUCH Ugerlmys antiquus MARTINSBRUCKE
TAVANNES
Llgerlmys Pseudotheridomys Psoudotherldomys+ llgorimys
BIERKELLER Eucricetodon nov.sp. SAFENWIL
Cervidae Eucricetodon Eucricetodon + Cervidae
BRUNNGRABEN
BRUTTELEN 2 Eucricetodon
ct. infralacto'rensis
Eucricetodon Lagopsls
MT. VULLY 1 aft. aquitantcus
?TEUFEN
Prolagus
It may come as a surprise that something can be made of such meager

material. However, it has to be considered that the evolution of the most important
mammal groups is quite well known from other parts of Europe. Therefore, it is not
difficult to date these relatively poor faunas.
Of course, not all small mammals are equally good for biochronology. In the
range of the Upper Marine Molasse the rodent families of the cricetids and eomyids
aze especially useful. Both families undergo a fast evolution in this time interval. In
addition, especially in the cricetids, new forms appear, from which no immediate
ancestors are known in the area of the Molasse. Such newcomers are considered
immigrants, and such immigrations are very useful for biochronology.
The following is a short characterization of each zone. The zone of Bruttelen Z

is based on a little fauna of small mammals discovered in 1986 by Jlirg Jost, a shark
teeth collector from Zofingen. Fossil mammals from Briittelen are known since the
end of the last century (see Studer, 1896; Stehlin, 1914), but the fauna consisted only
of large mammals. A new locality which we call Briittelen Z is only a few hundred
meters north of Briittelen 1. Among the small mammal remains of Briittelen Z are
molars of an Eucricetodon which I cannot distinguish (so far) from E. infralactorensis
on the basis of size and morphology. But I must point out that the assignment to
mammal zone MN 3a is not based only on the evolutionary level of this cricetid, but
also on the lithostratigraphical position of the locality. This locality is situated in a
profile only about 15m above a mammal site in a marl with freshwater molluscs, and
the age of this fauna is considered as Uppermost Aquitanian. The mammals from
Briittelen z, on the other hand, come from a marine sand with shark teeth. Since we
correlate the base of the Upper Marine Molasse with the Burdigalian transgression, the
fauna of Briittelen Z has been integrated into the mammal zone MN 3a.
The next younger zone of Bierkeller is mainly defined by a new species of

Eucricetodon. This species, which I intend to describe in the near future, is morpho-
logically very close to E. infralactorensis, but distinctly larger. Besides, we find in
this zone for the first time true cervids and Ligerimys. In the next older zone, that of
Briittelen Z, no Ligerimys has been found, but Pseudotheridomys is present.
178
Table z. Correlation chart.
Reference faunas Mediterranean

Correlations outside Lithostratigraphical
of the Assemblage Subdivision Stages
the Molasse (Steininger et a\.l
Zones
OSM
HIRSCHTHAL
MN 4 b VIEU/~~~~~~GES
~?.s:-
WATTWIL MN 4a LA ROMIEU ST.GALLEN
FORMATION
HINTERSTEINBRUCH MN 3b -----zOMM- BURDIGALIAN
LUZERNER
BIERKELLER
FORMATION
"'ZZ_,_
BR0TTELEN 2 MN 3a ESTREPOUY
MT. VULLY 1 MN 2b LAUGNAC AQUITANIAN

USM
From the locality of Bierkeller about 100 mammal teeth have been found so
far. This is very extraordinary, because in this locality we are dealing with a marine
sand, and the mammals are a minor by-product. All were found by the same Jiirg Jost
who washed large quantities of sediment to collect small shark teeth. The supposition
of a younger age for this Bierkeller locality compared with that of Briittelen Z results
also from its stratigraphic position. It is not situated in the same profile, I emphasize,
but is distinctly above the base of the Upper Marine Molasse.
The location of the reference fauna for the next zone, that of Hintersteinbruch,
is again distinctly higher in the series of the Upper Marine Molasse. This fauna, which
has yielded Ligerimys antiquus and Prolagus cf, vasconiensis, comes from freshwater
marls very near the coast. It is very conspicuous that cricetids are completely lacking
in this fauna.
The calibration of the fauna of Wattwil (see Frei, 1979), which I assign as refer-
ence fauna for the uppermost zone of the Upper Marine Molasse, was rather diffi-
cult. The characteristic mammal of this zone is a very small new species of
Ligerimys, which is of little use for biochronology. However, in this zone we find the
first modern cricetids, such as Megacricetodon and Eumyarion, the first elephants, and
Melissiodon is still present. On the basis of these first records I have correlated the
zone of Wattwil with the mammal zone MN 4a (level of La Romieu). From the litho-
stratigraphy we get little information for the datation of the Wattwil fauna, because
this locality is situated in eastern Switzerland, in a large series of freshwater sedi-
ments, a time equivalent of the Upper Marine Molasse.
The zone above that of Wattwil belongs to the Upper Freshwater Molasse. The
locality of Hirschthal, which I assign as reference fauna, was discovered by Job.
Hiirzeler in 19Z8 (see Hiirzeler, 193Z). New collections in 1981 yielded Ligerimys
florancei, Megacricetodon cf, bourgeoisi, and Anomalomys minor. These taxa place
this zone in the mammal zone MN 4b, the level of Su~vres and Vieux Collonges. Also
from its stratigraphic position we can conclude that this fauna must be very young. It
is situated in the so-called "Siis~brackwassermolasse," on top of a thick series of Upper
Marine Molasse. In addition to some isolated oysters, we find in this locality mainly
freshwater and land molluscs, and many leaves of land plants.
179
Table 3. Correlation of the lithostratigraphical units of
the Molasse with the international stages
(compiled by Beat Keller, Bern).
MERIAN (1836) BAUMBERGER INTE'RNATIONAL STAGES

STUDER (1839) (1934) (BERGGREN et al. 1985)
UPPER FRESHWATER Sarmatien ? TORTONI AN

SERRAVALLIAN
MOLASSE 'l/1/11111
Tortoni en 1//1111111
LANGHIAN
UPPER MARINE Helvetien BURDIGALIAN

?
MOLASSE Bureligal1en
111111111
LOWER FRESH- AQuitanien IIIII/III
AQUITANIAN
WATER MOLASSE Chattien CHATTIAN
Jll/111111
LOWER MARINE
MOLASSE Rupelien RUPELIAN
This zonation is only a beginning. Many of the mentioned faunas are still very
small, and also the lithostratigraphical relations have to be worked out. There is,
above all, the uncertainty that the base of the Upper Marine Molasse is diachronous in
different parts of Switzerland. At this moment several geological studies are under
way, whose aim, among other things, is to reconstruct the transgression of the Upper
Marine Molasse. With these studies we are anticipating for the near future, along with
more and more complete mammal faunas, to work out a more solid zonation of the
Upper Marine Molasse.
REFERENCES
Engesser, B. and Mayo, N., 1987. A biozonation of the Lower Freshwater Molasse
(Oligocene and Agenian) of Switzerland and Savoy on the basis of fossil
mammals. Miinchner Geowiss. Abh. (A), v. 10, p. 67-84.
Frei, H.-P., 1979. Stratigraphische Untersuchungen in der subalpinen Molasse der
Nordost-Schweiz zwischen Wagitaler Aa und Urnasch. Thesis of the Philosoph-
ical Faculty ll of the Ziirich University, Gotthard, Zurich, Z19 p.
Hiir:~;eler, J ., 193Z. Die Helvetien-Tortonien-Grenze im aargauischen Mittelland.
Ecolage geol. Helv., ZS, v. z, p. Z66-Z69.
Rutsch, R.F. and Schllichter, C., 1973. Stratigraphische Gliederung der Molasse im
bernischen Mittelland.,.Mitt. Naturf. Ges. Bern, N.F., v. 30, p. 85-90.
Stehlin, H.G., 1914. Ubersicht iiber die Saugetiere der schweizerischen
Molasseformation, ihre Fundorte und ihre stratigraphische Verbreitung. Verb.
Naturf. Ges. Basel, v. ZS, p. 179-ZOZ.
Studer, Th., 1896. Die Saugetierreste aus den marinen Molasseablagerungen von
Bruttelen. Abh. Schweiz. Palaont. Ges., ZZ, 4 7 p.
180
THE FAUNAL SUCCESSION OF THE BAVARIAN MOLASSE
RECONSIDERED- CORRELATION OF THE MN 5 AND MN 6 FAUNAS
Kurt Heiflig
Bayerische Staatssammlung fiir Palaontologie

und historische Geologie
Richard-Wagner-Str. 10/11
D-8000 Miinchen Z, Germany
INTRODUCTION
When in 1984 an intensive exploration of the exposures of the Upper Freshwater

Molasse started, there existed two biostratigraphical scales more or less independent
from one another. The first was the subdivision by Dehm (1951, 195~} in three units on
the basis of the occurrence of different proboscidean species, the "Altere Serie" with
only the small Gomphotherium angustidens (Cuvier), the "Mittlere Serie" with the
presence of both G. angustidens and Deinotherium bavaricum v. Meyer, and the
"Jiingere Serie" or "Hangendserie" with big species of both genera. Several local litho-
stratigraphical units have been assigned to these only paleontologically defined
"series," namely in the east of the Molasse basin.
The second scale was the succession of small mammal faunas based in 1964 by
Fahlbusch on the evolutionary changes of different hamster lineages. Most of the
more important sites were situated in the uniform sandy facies of the western part of
the basin and allowed no correlation with lithostratigraphical units. The advantage of
this scale was the correlation with the well known faunas of westem Europe. For bio-
facies reasons there was no correlation of large and small mammal sites. The only
rich fauna yielding both, Sandelzhausen, was not yet excavated sufficiently in the time
of Fahlbusch's work.
A new effort has been brought forward in 1984, when the dinosaur discussion le4
to the question (especially asked by W. Clemens) whether the Ries impact had caused
any extinctions, and if it is possible to recognize them in the faunas of the Molasse
basin. Nearly at the same time the stratigraphical difference between the "boulder
horizon" ("Brockhorizont"), originating from the Ries and the bentonite deposits was
emphasized by Scheuenpflug (1980). These bentonites are now recognized by Harr
(1976) as weathered andesitic volcanic ashes of unknown origin and therefore excluded
from the Ries phenomena. Nevertheless, they form a reliable time marker within the
puzzling fluviatile sediments of the Upper Freshwater Molasse. In the middle and
eastem part of the basin they have been recognized as the top of the "Mittlere
Serie." The debris of the Ries impact, however, also a reliable time marker, occur up
to 30 m deeper in the sequence, as it was recognized by Scheuenpflug (1980).
So the first explorations concentrated around the known occurrences of Ries

debris within molasse sediments and on sections below bentonite deposits. The first

00
II.)
... ....
/
/
--- ' \
I I
I
Goldberg• RIES • j
Adelschl ag
• •Eitensheim
''51
...... einberg
,. /___
•Stei nh eim •Gisse ltshausen
N
,,e \e /
•• "<!'-\\. /
/ I
/ 0 10 km 20 30
Fig. 1. Map of the mammal sites in the Upper Freshwater Molasse of Bavaria. Hatched = area of known outcrops
of the boulder horizon.
discoveries of small mammal faunas in 1984 led then to the investigation of a
combined section in the sand quarries of Gallenbach and the clay quarries of
Laimering and Unterneul, that was elaborated by Fiest (1986, unpublished). At the
same time geological mapping by Doppler (Bayerisches Geologisches Landesamt) in the
western part of the Bavarian Molasse yielded several new localities in deeper parts of
the Upper Freshwater Molasse, which had to be studied and which added also new
information (figure 1).
THE FAUNAS
Already the first faunas that have been discovered in the stratigraphic interval
between the boulder horizon and the top of the "Mittlere Serie" and even the fauna of
Unterneul just below the boulder horizon exhibited differences to any known small
mammal fauna of the Upper Freshwater Molasse. These faunas are characterized by:
1. The presence of the genus Cricetodon (figure Z) smaller than C. hagni Fahlbusch
but also different from C. sansaniensis Lartet.
z. The occurrence of a typical Megacricetodon minor (Lartet) instead of M. aff.

minor, the common small species of this genus in Bavaria (figure 3).
3. A further size increase of the bigger species of the most common cricetid genera,
Eumyarion, Megacricetodon, and Democricetodon (figures 4 and 5), compared with
the faunas assigned by Fahlbusch (1964) to the "Mittlere Serie."
These features show that all the faunas descr!ped by Fahlbusch except those
from the "Hangendserie" have to be placed into the "Altere Serie" (figure 6), whereas
the small mammal fauna of the "Mittlere Serie" was unknown up to now. Some other
data of these faunas are of some interest. First, there is the exact stratigraphical
position of the three specimens of Pliopithecus anti~uus Gervais in Bavaria. They are
all situated in or just above the "boulder horizon." econd, there is the persistence of
the opossum-like Amphiperatherium rather high in the "Mittlere Serie." Third, it may
be noted that a lot of taxa represented in the freshwater limestones of the Ries crater
lake are not yet found in the Molasse basin, especially the latest Cainotherium of
southern Germany.
THEFAUNALSUCC~ON
Fahlbusch based his succession on several lineages with a marked size increase.
Out of these the species group with the quickest change was chosen for a first
arrangement of the sites: Megacricetodon bavaricus/germanicus (figure 7). Fahlbusch
already noted the differences between the typical M. bavaricus from Langenmoosen
and the somewhat bigger M. aff. bavaricus fromthe later sites Rol3haupten and
a
Jettingen, which underwent further size increase in Oggenhof. Wu (198Z) found out
that the first increase took place rather rapidly, because the site of Puttenhausen,
placed in MN 5, yielded already M. aff. bavaricus. The name M. germanicus, given by
Aguilar (1980) to the much younger form (with equal size and equal morphology) from
Anwil (MN 8), was thought to be applicable also to this form, but there are some
doubts. So it is named here provisionally M. aff. germanicus.
New sites, comparable to the cited localities, have added some information to
this succession. In Bellenberg, in a rather deep level in the Upper Freshwater Molasse,
there occurs the typical M. bavaricus. In a distance of about 3 km and about 30 m
above, the site Betlinshausen has yielded mainly M. aff. germanicus but besides it a
few smaller individuals, that could be the latest representatives of M. bavaricus. Both
sites are separated by an erosional surface where the sand of Betlinshausen is over-
lying the silty marls of Bellenberg. So it remains uncertain if the sudden appearance
of M. aff. germanicus may be the result of a gradual size increase, masked by the
sedimentary hiatus, or if it is a real event, as suggested by the probable common
occurrence of both species.
183
8
mm
2,4
,•'
,'
,'
2,3 v
.-·-· -·-·-.
2,2
:
: /~;,..Jl- .. -- ..
. .
....,.-~·=- ~---=-·~. . . .---::~.\--............. 0
: ~'/--·"
•_.-/ .. ' \b . ·\' \
1
.r·.. / V ... ·, ! ;'
2,1 I I
-'-' ' II V II
... I / /
/;/
//
.-'/ ('· ... V
J' '• ......;.;.. .. ··· j/
1//·
..
,..... /
2,0
/
/
/ V V •II .-·-;:;?._......
'--.L.-·-·-·--~-·-v ·-·-·-..;-;:. ""/"'
1,9
(ov
' '....._______
v 0
--
·.'-.-.. ...-··--~..---'
...........
1,8 0
"
2,7 2,8 2,9 3,0 3,1 3,2 3,3 3,4 3,5 3,6 L, mm
8
, .. ·.
mm
.·.---·-·- ·-·-
J/,·,':· .. - \
,/ , II i
1,9 I
.
/
II .·,
. II
/
/
1,8
,/·~--..::.----- ..... II II
/
v_~,..- /m '\ 11
/
~ : V II I
:V __,
II
II I /
/.-'
1,7 VII / ,'
. -'l
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.
.- ....-·
/
v ..... ·..../ /
/
/
...
;......-: .......... v
/
0 •••
1,5
1,4
2,4 2,5 2,6 2,7 2,8 2.9 3,0 3,1 L, mm

2,2 2,3
0 Ebershausen II Laimering 2 - - meini, Vieux Co!longes
Cricetodon
o Unterneul 0 Unterzolling - - jotae, Manchones
G 8ubenhausen II Hall Steinberg
v Galien bach 2 b _ .. _ Goldberg
" Gallenbach 2c · · · · · hagni, Anwil

·
Fig. z. Scatter diagram of the first upper and lower molars of different
species and samples of Cricetodon . L = length, B = width, both
in mm.
184
8
mm
.······.
..-· - . , --_,~--, .i
/ _;..y,....,...::...----....... .\
/-:'/·~·'\ ")-.'' \
1,0
// _.· \
// ,. . \ I
/
/• /~~ / ! ......'!1···· J
I / .,.;_ I «1-_...
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I · 11 v; e/ /
//
./
0.9 i • / /
v/ __ _.._.,.,. /
\ \
-·____
.
...../-·-·
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..,__, l"ff \... v
·.. ..- ..
/
·......_"-..- ........._ /
o.s
1,2 1,3 1,4 1,5 1,6 L. mm
8
mm
/.···· -----
~·_.,~;:;\
...-:::... -- .
--- \ :\ :"'.
///-----··-· ·-··ax·. i
/ / .;<-·· ...--;) i
0,9
. '.
j--:7· ! v........ J/ l
\ ...···A/
)· v:···•. . . . . ! ...···:::~/
/ I ~.,.
. '\, • ----.::?'·· /
0,8 . / e e ~ -----::-......-· /
'-.. I .v- ~--
.--- -:...~~.:.::!. ....-
0,7
1,1 1,2 1,3 I,C 1,5 L, mn

Megacricetodon minor u. similis II Laimering 2 M. aft. minor, Sande!zhausen
• Steinheim M. minor, Goldberg
t> Unterneu! M.minor, Steinberg
• Gisse!tshausen M. minor, Sans an
e Ziemetshausen M. simi lis, ·~ungere Serie"
V Ga!!enbach 2b
"f' Gallen bach 2 c
Fig. 3. Scatter diagram of the first upper and lower molars of the
small Megacricetodon species of Bavaria, compared with M.
minor from Sansan. L -length, B =width, both in mm.
185
8
mm
1,6
1.5
1,4
1,3
1,2
1,9 2,0 2,1 2,2 2,3 2,4 L, mm
8
mm
1,4
v
1,3
1,2
1,1
1,0 '... ..... _______ _ --
1,7 1,8 1,9 2,0 2,1 2,2 L, mm

Eumyarion bifidus
Puttenhausen I:. Sa II mannsberg
Sandelzhausen o Ebershausen
Gisseltshausen 1a • Affalterbach
Gis seltshausen1b V Gallenbach 2b
D Unterzolling
• Steinheim
Fig. 4. Scatter diagram of the first upper and lower

molars of Eumyarion bifidus from various sites
in Bavaria. L =length, B =width, both in mm.
186
B 0
mm 0
0
1,5
1,8 2,3 2,4 L,mm
B
0
mm
1,4
1,3
1,2
I
1,1
"'" ..
.-"' "' .. .··
'·.'"'--T.~:-: ,."""
....... .....
1,6 1,8 1,9 2,0 2,1 2,2 2,3 L,mm

Megacricetodon bavaricus Megacricetodon germanicus M.sp.
--- Langenmoosen Anwil 0 Ebershausen
Puttenhausen
Oggenhof
Affalterbach
e Gisseltshausen
Fig. 5. Scatter diagram of the first upper and lower molars of big
Megacricetodon species from Bavaria, compared with M.
germanicus from the type locality (Anwil). L = length, B =
width, both in mm.
187
Fahlbusch 1964 Hei~ig 1988
:Harktl :
:Giggenhausen Giggenhausen : Anwil

: Kleineisenbach
: :
:oggonbo'\
La Grive :?Hader Steinheim
:
: La Grive ?
:
: Laimering 2* : Manchones?
: Gallenbach 2*:
:Jettingen Unterzolling*:
~'c
:Ro~haupten Unterneul* :
Sans an : Ebershausen* : Sans an
Hanchones: ~ffalterbach*:
: Oggenhof :Collet Redon
: Puttenhausen :
: Jettingen :
: Betlinshausen:
: Ro~haupten :
:
:Langenmoosen Langenmoosen :
: Bellenberg :
Fig. 6. The correlation of the micromammal sites in the Upper

Freshwater Molasse, reinterpreted in the light of the
new localities of the "Middle Series" and just below (*).
The next step in the size increase, that was found in the site Oggenhof by
Fahlbusch, is represented also in the new locality Affalterbach near Pfaffenhofen.
For geological reasons this site is thought to be situated deeper than the boulder
horizons in this region. Also the new locality of Ebershausen, which has yielded the
first Cricetodon lies below this time marker. There has been found the (until now)
biggest species of Megacricetodon, that is named here as M. sp. due to insufficient
documentation.
Later than this fauna, that may represent the base of the "Mittlere Serie," the
record of Megacricetodon is so scarce that this genus is of no stratigraphical value.
So the genus Cricetodon may be used within this unit. The first occurrence of this
genus is assumed to be contemporaneous with the reimmigration of Deinotherium,
probably caused by the same ecological change. The specimens from the localities
Ebershausen, Bubenhausen, and Unterneul are of small size, comparable to Cricetodon
meini Freudenthal from Vieux Collonges. The later locality Gallenbach 2, about 15 m
above the boulder horizon, has yielded bigger specimens comparable to the size of
Cricetodon jotae Freudenthal, but of rather different morphology. This species
continues its size increase until the top of the "Mittlere Serie" at the site of
Laimering 2, just below a bentonite deposit.
In the same sites there occurs also the smaller lineage of Megacricetodon, that
begins in Unterneul with M. minor (instead of M. aff. minor) and comes near the size
of M. similis Fahlbusch in Laimering Z and in thesediments of the Ries crater lake.
188
Yestern Europe Middle Europe
locality I length of M1 13 14 15 16 17 18 19 20 21 22113 14 15 16 17 18 19 20 21 22 23 24 (0,1 mm) I locality
Nombrevilla ibericus I
I germanicus Anwil
I I
I I
La Grive gregarius I I I
/ >» >>>:» >>» >» I»>>»»>>-. ...!..--- I gregarius Steinheim
I
crusafonti ' I I
!1anchones I
II ~~ I
I ,~ I
~~
I
gersi I - - ,~ I
Sans an I I
I I 1
.................. ·
H. sp. Ebershausen • .•••.••
I I
Collet Redon germanicus I I <« « « « <« « (', Affalterbach
I I
__ - '···crff • germanicus Oggenhof
I I ««<«« -----;:'
I I ,... ', Gisseltshause n
I I
I I ''
,-----; Puttenhausen
I I
I I
{ . - ............ Jettingen
I I
I I ,' --- 1 ----,. Betlinshausen
I I
I I I o Ro~haupten
I --,--
I I
I I .__j bavaricus Langenmoosen
\
I I ,/
I I Bellenberg
I I ............ ... .........
I I Franzensbad
I I
I
Las Planas IV collongensis ~
Forsthart
I
,
__ ' Rauscherod
La Romieu
Fig. 7. Size evolution (of M/1) in the bigger species of Megacricetodon, M. collongensis, M. gregarius, M. bavaricus, M.
germanicus, M. ibericus, and M. sp. in the light of the new populations. >>>>>::.. and <<<<<< = migrations.
Cll
co
CORRELATIONS
The only sites of comparable age outside the Molasse basin in southern Germany
are Steinheim a.A. and the sites in the freshwater limestones of the Ries crater
Goldberg and Steinberg. The latter two can be easily correlated with the latest fauna
of the "Mittlere Serie," Laimering Z with at least three common species of cricetids:
Cricetodon sp., Megacricetodon minor/similis and Democricetodon gracilis
Fahlbusch. The two first show nearly no difference in size and really none in morphol-
ogy between the sites of the Molasse basin and the Ries crater. The third one exhibits
no changes during the observed time interval. Democricetodon aff. gaillardi in the
Ries sediments is still morphologically similar to the tnore primitive D. mutilus
Fahlbusch in the sites of the "Mittlere Serie," but different in size. So this species
cannot be used for direct correlation.
The few remains of small mammals of the Steinheim basin cannot be compared
with the species in the other sites of southern Germany. The only common species
Democricetodon aff. gaillardi is not yet studied in detail. The species Megacricetodon
gregarius is unknown elsewhere in Germany and is probably an immigrant from
western Europe.
The correlation with faunas in western Europe is rather difficult due to the
presence of endemic lineages in both regions. Cricetodon starts with a size and
morphology similar to C. meini from Vieux Collonges, but probably at a considerable
later time. Its lineageshows an evolution that is different in southern Germany from
France and Spain and which leads directly to the later endemic species Cricetodon
hagni Fahlbusch in the "Ji.ingere Serie." The phylogeny of Eumyarion is not yet clearly
understood. The genus shows in its bigger species a size increase that comes near to
the size of Eumyarion medius from Sansan, but it exhibits the morphology of the
different lineage of E. bifidus Fahlbusch, that is confined to southern Germany.
Megacricetodon is split up early in two lineages or more: in France and Spain there
occurs the M. collongensis-crusafonti-gregarius lineage, in Germany and Switzerland
the M. bavaricus/germanicus lineage. Both show several complications. There are
two connections. There is one occurrence of M. germanicus in France, east of the
Rhone at the site of Collet Redon. But the Size of this species probably remains
unchanged during the time from Oggenhof (probably MN 5) up to Anwil (MN 8). The
immigration of M. gregarius in Steinheim gives a clear time marker and confirms the
position of this site within the MN 7 "zone." Another immigration is the "sudden"
occurrence of Megacricetodon minor at the base of the "Mittlere Serie." This species
shows a size increase and can therefore be correlated by its similar size with the
famous site of Sansan, whereas the later localities show the evolution of this species
toward M. similis. So the only good correlation can be made at the base of the
"MittlereSerie" with Sansan.
SUMMARY
New small mammal sites, found in connection with the boulder horizon (Ries
debris within the molasse) and the volcanic bentonites, represent the fauna of the
"Mittlere Serie" of Dehm. This fauna is different in the Cricetids from any hitherto
known fauna in the molasse basin. So all the faunas formerly ascribed to this series
have to be placed into the "Altere Serie." The faunal association of the "Mittlere
Serie" begins at a level below the boulder horizon with the immigrations of Cricetodon
and Megacricetodon minor, thought to be contemporaneous ,~:Vith the Deinotherium
immigration. It is separated by an erosional hiatus from the "Altere Serie." This can
be divided into three parts based on the size of the Megacricetodon bavaricus/
germanicus lineage. The lower part of the "Mittlere Serie" can be correlated with
Sansan, whereas its higher parts, contemporaneous with the crater sediments of the
Ries, are younger. Steinheim in the neighboring smaller crater basin is even more
recent. Its correlation with La Grive is confirmed by the immigration of Mega-
cricetodon gregarius.
190
ACKNOWLEDGMENTS
I am grateful to Prof. Dr. W.v. Engelhardt, L. Scheuenpflug, and Dipl. Geol. K.

Ulbig for the communication of new outcrops of the boulder horizon. I want to thank
G. Berger for the communication of new small mammal sites and P. Veit for his
assistance in fieldwork and the technical processing of samples. To K. Dossow I am
indebted for the preparation of the scatter diagrams.
I want to express my deepest gratitude to my friend and colleague Prof. Dr.

Fahlbusch for a lot of detailed discussions and his patience with my premature ideas
on his beloved hamsters.
REFERENCES
Aguilar, J.-P., 1980. Nouvelle interpretation de !'evolution du genre Megacricetodon

au cours du Miocene. Palaeovertebrata Mem. Jub. R. Lavocat, p. 335-364.
a
Aguilar, J.-P. and Clauzon, G., 1979. Un. gisement mammiferes dans la formation
lacustre d'age Miocene moyen du Collet Redon pres de St-Cannat (Bouches du
Rhone), Implications stratigraphiques. Palaeovert., v. 8(5), p. 3Z7-341.
Aktas, A., 1987. Altquartare Schotter der Zusam-Platte, Bayerisch Schwaben.
Sonderver. Geol. Ins. Univ. Koln 6Z, p. 1-100.
Cicha, I., Fahlbusch, V., and Fejfar, 0., 197Z. Die biostratigraphische Korrelation
einiger Jungtertiirer Wirbeltierfaunen Mitteleuropas. N. Jb. Geol. Pal. Abh, v.
140(Z), P• 1Z9-145.
Dehm, R., 1951. Zur Gliederung der jungtertiaren Molasse in Suddeutschland nach
Saugetieren. N. Jb. Geol. Palaont. Mh. 1951, p. 140-15Z.
Dehm, R., 1955. Die Saugetier-Faunen in der Oberen Siit3wassermolasse und ihre
Bedeutung (ur die Gliederung, in Erl. geol. Ubers-Kte. sUddt. Molasse, ed. Bayer.
Geol. L.A., p. 81-88. -
Engesser, B., 197Z. Die obermiozane S"augetierfauna von Anwil (Baselland).
Tatigkeitsber. Naturforsch. Ges. Baselland Z8, p. 35-363.
Fahlbusch, V., 1964. Die Cricetiden (Mamm.) der Oberen S\iawasser-Molasse Bayerns.
Abh. Bayer. Akad. Wiss, Math.-Naturw. Kl. N. F. 118, p. 1-136.
raphy of the European Tertiary. Newsl. Stratigr., v. 5, p. 16Q-167.
Fahlbusch, V., 1981. Miozan und Pliozan - Was ist das? Zur Gliederung des
Jungtertiars in Siiddeutschland. Mitt. Bayer. Staatsslg. Pafaont. hist. Geol., v.
Z1, p. 1Z1-1Z7.
Fejfar, 0., 1974. Die Eomyiden und Cricetiden (Rodentia, Mammalia) des Miozans der
Tschechoslowakei. Palaeontographica A 146, p. 100-180.
Fiest, W., 1986. Lithostratigraphie und Schwermineralgehalt der Oberen
Siiflwassermolasse im Bereich um die Gallenbacher Mulldeponien zwischen
Aichach und Dasing. Unpubl. Diplome Thesis, Munich, 119 p.
Harr, K., 1976. Mineralogisch-petrographische Untersuchungen an Bentoniten in der
siiddeutschen Molasse. Unpubl. Thesis.
Heil3ig, K., 1986. No effect of the Ries impact event on the local mammal fauna.
Modern Geology, v. 10, p. 171-179.
Heieig, K., 1989. Neue Ergebnisse zur Stratigraphie der Mittleren Serie der Oberen
Siiawassermolasse Bayerns. Geol. Bavarica, v. 93 (in print).
Heiaig, K. and Fiest, W., 1987. Neue Funde von Pliopithecus in Bayern. Mitt. Bayer.
Staatsslg. Palaont. hist. Geol., v. Z7, p. 95-103.
Heizmann, E.P.J. and Fahlbusch, V., 19&~· Die mittelmioziine Wirbeltierfauna vom
Steinberg (Nordlinger Ries), Eine Ubersicht. Mitt. Bayer. Staatsslg. Palaont.
hist. Geol., v. Z3, p. 83-93.
Hofmann, F., 1965. Die stratigraphische Bedeutung der Bentonite und Tufflagen im
Molassebecken. Jber. Mitt. oberrh. geol. Ver. N. F., v. 47, p. 79-90.
Mein, P., 1975. R~sultats du groupe de travail des Ve~teb.res, in Report on Activity of
the R.C.M.N.S. Working Group (1971-1975), p. 78-81. -
Mein, P. and Freudenthal, M., 1971. Les Cricetidae (Mammalia, Rodentia) du NeogEme
moyen de Vieux Collonges, Prt. 1: Le genre Cricetodon Lartet 1951. Scripta
Geol., v. 5, P• 1-38.
191
Scheuenpflug, L., 1980. Neue Funde ortsfremder Weiajuragesteine in Horizon ten der
siidbayerischen miozanen Oberen Siif3wassermolasse um Augsburg. Jber. Mitt.
oberrh. geol. Ver. N. F., v. 62, p. 131-142.
Stephan, W., 1952. Ein tortoner vulkanischer Brockhorizont in der Oberen
Siif3wassermolasse Bayerns. Geol. Bav., v. 14, p. 6-85.
Wu, W., 1982. Die Cricetiden (Mammalia, Rodentia) aus der Oberen Siiawasser-
Molasse von Puttenhausen (Niederbayern). Zitteliana, v. 9, p. 37-80.
Ziegler, R. and Fahlbusch, V., 1986. Kleinsliuger-Faunen aus der basalen Oberen
Siitivasser-Molasse Niederbayems. Zitteliana, v. 14, p. 3-58.
192
STRA'nGRAPHY OF NEOGENE MAMMALS OF POLAND
Kazimierz Kowalski
Institute of Systematic and Experimental Zoology

Polish Academy of Sciences
31-016 Krak6w
Stawkowska 17, Poland
INTRODUCTION
First information concerning Neogene mammals within the present territory of

Poland was gathered in Silesia, at that time a part of Prussia, in the second half of the
19th century. In 1859, R. Hensel described as a new species Prox (now Euprox)
furcatus, a primitive deer, from the vicinity of Gliwice. In 1888, E. Koken reported
another discovery of Miocene mammals from the same region. A much richer fauna
was described from middle Miocene sediments near' Opole (Oppeln) in Silesia by R.
Wagner in 1913.
Data from other regions of Poland began to accumulate much later. In 1934, a
Polish geologist, J. Samsonowicz, discovered a Pliocene bone breccia in W~ze in
central Poland. Its paleontological study began in 1950, when J. Stach described
several new species of carnivores from it. In the sixties the number of known fossil
localities began to grow and studies of different groups of mammals, particularly
small ones, were undertaken by various scholars. The research concentrated in the
Institute of Systematic and Experimental Zoology of the Polish Academy of Sciences
in Krakow, where the largest part of collections was deposited. The Institute of
Paleobiology of the Polish Academy of Sciences in Warsaw and the Department of
Paleozoology of the Wroclaw University were also active in this study.
MIOCENE
The oldest mammalian remains in Poland were found in middle Miocene layers
corresponding to the zones MN 4 through MN 6. They were discovered in Be1chatow
(BEl), situated halfway between Cracow and Warsaw, in a mine of lignite. Strip-
mining discovered there extensive layers of lignite with interclating layers of limnic
chalk mainly composed of shells of molluscs, including both aquatic and terrestrial
snails. The molluscan fauna with numerous Oriental elements suggests a tropical
climate and a forest environment. Mammal fauna recently discovered in chalk
contains undetermined insectivores, carnivores, and lagomorphs, and numerous rodent
teeth. They belong to species from the families Sciuridae, Eomyidae, Gliridae,
Castoridae, and Cricetidae, and have not yet been studied in detail. Presence of the
genera Neocometes and Anomalomys is especially noted. Only a small part of the
rodent material was identified by V. Fahlbusch; in the paper by G!azek and
Szynkiewicz (1987), Eumyarion cf. weinfurteri, Democricetodon cf. vindobonensis, and
D. gracilis were listed from this locality on the basis of his determinations (table 1).

co
.:.
KIELCE
..,
---------...'-11-·~ .PR ~KD
,.,
·--.
'·\.,.,.
•
- ...e.
a 'ID sa 'IDD km
~
Fig. 1. Neogene mammal localities in Poland. 1 = Miocene; Z = Pliocene. BE = BeJchatow; KA = Kamyk;

KD = Kadzielnia; KI = Kielniki; PO= Podlesice; PR = Przeworno; RK = R~bielice Krolewskie; WE= Wl(ze.
Table I. Mammalian species from the Neogene of Poland. Locality abbreviations: PRI = Przeworno I; PRZ = Przeworno Z; OPI =
Opole I; OPZ = Opole Z; BEZ = Be1chatow Z; PO= Podlesice; WEI = W~ze I; RKI = R~bielice Krolewskie I; RKZ = R~bielice
Krolewskie z. M = species known only from the Miocene localities other than those listed before; P = species known only from
the Pliocene localities other than those listed before.
Localities
Species PRI PRZ OPI OPZ BEZ PO WEI RKI RKZ M P
Jnsectivora
Metacordylodon schlosseri Andreae, I904 X
"Erinaceus sansaniensis Lartet, I95I" X
Erinaceus samsonowiczi Sulimski, I959 - X
Scaptonyx(?) dolichochir (Gaillard, I899) X X X
Scapanulus agrarius Skoczen, I980 X X
Neurotrichus(?) polonicus Skocze:ii., I980 - - X
Condylura kowalskii Skocze:ii., I976 X X
Condylura izabellae Skoczen, I976 - - - - - X
Talpa minuta Blainville, I838 X
Talpa minor Freudenberg, I9I4 X X X
Talpa fossilis Petenyi, I864 X
Desmana nehringi Kormos, I9I3 X X
Galemys sulimskii Riimke, I985 X
Galemys kormosi (Schreuder, I940) - X X
Sorex minutus Linnaeus, I766 X X
Sorex subminutus Sulimski, I96Z X
Sorex runtonensis Hinton, I911 X X X
Sorex praealpinus Heller, I930 X
Petenyia robusta Rzebik-Kowalska, 1988 X -
Petenyia hungarica Kormos, 1934 - X X X
Blarinella dubia (Bachmayer and Wilson, 1970) X -
Blarinella europaea Reumer, 1984 X X X
Zelceina soriculoides (Sulimski, 1959) X X
Deinsdorfia hibbardi (Sulimski, I96Z) X
Blarinoides mariae Sulimski, 19 59 X X X X
Mafia dehneli (Kowalski, 1956) X X
co
U1 Sulimskia kretzoii (Sulimski, 196Z) X
(cont.)
CD
Ol Table 1 (continued)
Localities
Species PRl PRZ OPl OPZ BEZ PO WEI RKl RKZ M p
Episoriculus gibberodon (Petenyi, 1864) - - - - - X X

Neomysorex alpinoides (Kowalski, 1956) - - - - - X
Beremendia fissidens (Petenyi, 1864) - - - - - - X X X
Beremendia minor Rzebik-Kowalska, 1976 - - - - - - - X
Amblycoptus topali Janossy, 197Z - - - - - - - - - - X
Paranourosorex gigas Rzebik-Kowalska, 1975 - - - - - X
Paenelimnoecus pannonicus Kormos, 1934 - - - - - - X
Chiroptera
Rhinolophus kowalskii Topal, 1979 - - - - - X - - X
Rhinolophus hanaki Woloszyn, 1988 - - - - - X
Rhinolophus wenzensis Wol'oszyn, 1988 - - - - - - X
Rhinolophus cf. variabilis Topal, 1975 - - - - - - X
Rhinolophus neglectus Heller, 1936 - - - X
Rhinolophus macrorhinus Topal, 1963 X
-
-- - -
-- -- .--
Miniopterus approximatus Wol'oszyn, 1988
-- - -
- - - - X
Plecotus rabederi WoJoszyn, 1988 - - - X X
--
Eptesicus kowalskii Wo1oszyn, 1988
-
- - - - - X
Eptesicus mossoczyi Wol'oszyn, 1988 - - - X
Myotis danutae Kowalski, 1956 X X
-- --
- - -
Myotis podlesicensis Kowalski, 1956 - - - - - X X
Myotis helleri Kowalski, 196Z - - - - - X
Myotis delicatus Heller, 1936 X
--
- - - -
Myotis gundersheimensis Heller, 1936 - - - - - X X
Myotis cf. exilis Heller, 1936 - - - - - X X
Primates
Pliopithecus antiquus (Blainville, 1839) - X X
Localities
Lagomorpha
Eurolagus fontannesi (Deperet, 1887) - - X
Ochotona polonica Sych, 1980 - - - - - - - - - - X
Hypolagus beremendensis (Petenyi, 1864) - - - - - - X X X
Rodentia
Sciuridae
Palaeosciurus cf. fissurae (Dehm, 1950) - - - X
Spermophilinus bredai (Meyer, 1848) - - - X X
Miopetaurista gaillardi (Mein, 1970) - - - X
Miopetaurista gibberosa (Hofmann, 1893) - - X X
Miopetaurista cf. thaleri (Mein, 1970) - - - - - - - X
Blackia polonica Black and Kowalski, 1974 - - - - - X X X X
Pliopetaurista dehneli (Sulimski, 1964) - - - - - X - X X
Pliopetaurista cf. pliocaenica (Deperet, 1897) - - - - - - - - - - X
Pliopetaurista schaubi (Sulimski, 1964) - - - - - - X
Pliopetes hungaricus Kretzoi, 1959 - - - - - X X
Sciurus warthae Sulimski, 1964 - - - - - X X X X
Tamias orlovi (Sulimski, 1964) - - - - - X X X X
Eomyidae
Keramidomys carpathicus (Schaub and Zapfe, 1953) - - - X
Keramidomys mohleri Engesser, 197Z - - - - - X
Estramomys cf. simplex Janossy, 1969 - - - - - - - X
Leptodontomys cf. catalaunicus (Hartenberger, 1967) - - - - - X
co (cont.)
--.1
CQ Table 1 (continued)
co
Localities
-
Castoridae
Chalicomys jaegeri Kaup, 183Z X - X X
Steneofiber eseri (Meyer, 1846) - - - X
Steneofiber minutus (Meyer, 1838) - - - X
Steneofiber w«:nzensis Sulimski, 1964 - - - - - - X
Cricetidae
Eumyarion bifidus (Fahlbusch, 1964) - - - X
Eumyarion cf. weinfurteri (Schaub and Zapfe, 1953) - - - - X
cf. Megacricetodon bavaricus Fahlbusch, 1964 - - - X
Democricetodon gaillardi (Schaub, 19Z5) - - X X
Democricetodon cf. vindobonensis (Schaub and Zapfe, 1953) - - - - X
Democricetodon gracilis (Fahlbusch, 1964) - - - - X
Kowalskia magna Fahlbusch, 1969 - - - - - X
Kowalskia polonica Fahlbusch, 1964 - - - - - X
Epimeriones austriacus Daxner-H'ock, 197Z - - - - - - - - - X
Epimeriones progressus Kowalski, 1974 - - - - -- X
Baranomys loczyi Kormos, 1933 - - - - - - - - - - X
Microtodon kowalskii (Kretzoi, 196Z) - - - - - X
Trilophomys pyrenaicus (Deperet, 1890) - - - - - - X
Anomalomys gaudryi Gaillard, 1900 - - - X
Arvicolidae
"Trilophomys" canterranensis Michaux, 1976 - - - - X
Prosomys insuliferus (Kowalski, 1958) - X
- - -- -
Germanomys weileri Heller, 1936 - - - - - - X
Stachomys trilobodon Kowalski, 1960 - - - - - - X X X
Dolomys cf. milleri Nehring, 1898 - - - - - X
Mimomys altenburgensis Rabeder, 1981 - - - -- - - - X X
Mimomys gracilis (Kretzoi, 1959) - - - - X
Mimomys hassiacus Heller, 1936 X
--
-
--
- - -
Localities
PRZ OP1 OPZ BEZ PO WE1 RK1 RKZ M p
Species PR1
- - - - - - - X X
Mimomys polonicus Kowalski, 1960 X
Mimomys cf. reidi Hinton, 1910 - - - - - - -
- - - - - - X
Mimomys cf. stehlini Kormos, 1931
- - - - - - X X
Propliomys hungaricus (Kormos, 1934)
- - - - - - - X X
Villanyia cf. exilis Kretzoi, 1956 X
- - - - - - - X
Synaptomys europaeus Kowalski, 1977
Spalacidae
- - - - - - X X
Prospalax priscus (Nehring, 1897)
Muridae
- - - - - - - X
Apodemus dominans Kretzoi, 1959
- - - - - - X X
Micromys cf. praeminutus Kretzoi, 1959
- - - - - X
Parapodemu s coronensis Schaub, 1938 - X
Parapodemu s lugdunensis Schaub, 1938 - - - - - - - - -
- - - - - - X
Parapodemu s schaubi Papp, 1947
- - - - - - X X
Rhagapodem us frequens Kretzoi, 1959
Gliridae
- - - X
Glis cf. vallesiensis Aguf!tf, 1981
- - - - - X X X
Glis minor Kowalski, 1956
- - - - X
Glirudinus sp.
- - - - - X
Glirulus pusillus (Heller, 1936)
- - - - - X X X
Muscardinus pliocaenicus Kowalski, 1963
- - - - - X
Muscardinus cf. dacicus Kormos, 1930
Seleviniidae
- - - - - - X
<D Plioselevini a gromovi Sulimski, 196Z
<D
(cont.)
N
0 Table 1 (continued)
0
Localities
---
Zapodidae
Sminthozapus intermedius Bachmayer and Wilson, 1970 - - - - X
Sminthozapus janossyi Sulimski, 196Z X X
-- -
- - - -
Hystricidae
Hystrix primigenia (Wagner, 1848) - - - - - - X
Carnivora
Pseudaelurus quadridentatus (Blainville, 1843) X
Pseudaelurus lorteti Gaillard, 1899 - X
Felis wenzensis Stach, 1961 - - - - - - X
Alopecocyon goeriachensis (Toula, 1884) - - X
Nyctereutes cf. donnezani (Deperet, 1890) - - - - - X
Ursavus brevirhinus (Hofmann, 1887) - - X
--
Ursavus primaevus (Gaillard, 1899)
- - - - - -
- - X
Agriotherium intermedium Stach, 1957 - - - - - - X
Ursus wenzensis Stach, 1959 - - - - - - X
Heterictis oppoliensis (Wagner, 1913) - - X
Trochotherium cyamoides Fraas, 1870 - - X
Martes wenzensis Stach, 1959 - - - - - - X
Mustela pliocaenica Stach, 1959 - - X
Baranogale helbingi Kormos, 1934 X X X
- --
-- -- --
Arctomeles pliocaenicus Stach, 1951 - - - - - - X
Perissodactyla
Tapirus cf. telleri Hofmann, 1893 - - - X
Aceratherium simorrense (Lartet, 1851) X X X
Aceratherium tetradactylum (Lartet, 1835) - - X
Brachypotherium brachypus (Lartet, 1884) X X X
Dicerorhinus megarhinus (Christol, 1835) - - - - - - X
Localities
PR1 PRZ OP1 OPZ BEZ PO WE1 RK1 RKZ M p
Species
Anchitherium aurelianense (Cuvier, 18Z5) X X X

Chalicotherium grande (Lartet, 1837) - - X
Proboscidae
Tetralophodon longirostris Kaup, 183Z - - - - - - - - - - X
Gomphotherium angustidens (Cuvier, 1806) X - X

Mammut praetypicum (Schlesinger, 1919) - - - - - - - - - - X
Artiodactyla
Conohyus simorrensis (Lartet, 1851) X X X
Taucanamo sansaniense (Lartet, 1851) - X
Dorcatherium crassum (Lartet, 1851) X X
Palaeomeryx eminens v. Meyer, 1847 - - X
Euprox furcatus (Hensel, 1859) X X X
Procapreolus wenzensis (Czyzewska, 1960) - - - - - - X
Cervus warthae (Czyzewska, 1968) - - - - - - X
Cervus cusanus Croizet et Jobert, 18Z8 - - - - - - - - - - X
Cervus pardinensis Croizet et Jobert, 18Z8 - - - - - - - - - - X

Arvernoceros ardei (Croizet et Jobert, 18Z8) - - - - - - - - - - X
Croizetoceros ramosus (Croizet et Jobert, 19Z8) - - - - - - - X
Unpublished data were kindly submitted by B. Rzebik-Kowalska, A. Nadachowski, V. Fahlbusch, M. Wolsan, T. Czyzewska.
1\o.)
0
Other Miocene localities, situated in southwestern Poland, in Silesia, are of
slightly younger age and can be correlated with zones MN 5-7 (figure Z). One of them
is Opole. The locality studied by R. Wagner and published by him in 1913 (OP1)
contained bones of vertebrates and shells of snails in loams filling a channel and
covered by lignites. This fauna was very rich in bones of larger mammals. Among
insectivores there was Metacordylodon schlosseri, among primates Pliopithecus
antiquus, among lagomorphs Eurolagus fontannesi. Only scarce remains of rodents
were described; they belonged to Miopetaurista gibberosa, Chalicomys jaegeri, and
Democricetodon cf. gaillardi (see Schaub, 19Z5), all known also from Opole z.
Opole 1
was the richest Polish Miocene fauna of carnivores and ungulates. Some determina-
tions need revision; unfortunately most of the material cannot be located. Among
perissodactyls there were Aceratherium simorrense and A. tetradactylum,
Brachypotherium brachypus, Anchitherium aurelianense, and Chalicotherium grande.
The artiodactyls from Opole 1 contain Conohyus ·simorrensis, Euprox furcatus, and
Palaeomeryx eminens. Finally, there was a mastodon, Gomphotherium angustidens.
EPOCHS
Romanian
PLIO
CENE Decian
late
Pontian
middle Sarrnatian
MID
CENE
Fig. z. Stratigraphic position of Neogene mammal

localities in Poland. BE Z = Be1chatow Z; KA =
Kamyk; KD = Kadzielnia; KI 3A = Kielniki 3A;
K1 3B = Kielniki 3B; OP 1 - Opole 1; OP Z =
Opole Z; PO = Podlesice; RK 1-Z = R~bielice
Krolewskie 1 and Z; WE 1 = W~ze 1.
202
In 1950, a new locality with fossil vertebrates was discovered in Opole (OPZ). It
is situated about 1 km from Opole 1; the bones were also found in loams which were,
in this place, not covered by lignites. According to some authors (G!azek and
Szynkiewicz, 1987), this new locality is slightly younger than Opole 1 and represents
biozone MN 7. The fauna contained a tooth of Tapirus cf. telleri (Ryziewicz, 1961)
and remains of 13 rodent species (Kowalski, 1967). Among them, of particular strati-
graphic importance, were Democricetodon gaillardi and D. gregarius bavaricus,
Eumyarion bifidus, and Anomalomys gaudryi. In my opinion there is no reason to
believe that both localities in Opole are of different age.
Other faunal assemblages of similar age are known from the locality Przeworno
in Lower Silesia. The fauna of Przeworno, discovered in 1970, is preserved in karst
fissures and channels in a quarry of marbles of undetermined age (probably Protero-
zoic). Some bones were found in loams filling horizontal channels at the bottom of the
quarry (Przeworno 1 - PR1) (G1azek et al., 1971). More fossil material was excavated
from a vertical fissure situated in the western wall of the quarry (Przeworno Z -
PRZ). It was some 1 Z m deep and was filled with clay and blocks of marble. It is
probable that during the accumulation of the fauna an open, vertical fissure func-
tioned as a trap for terrestrial animals. Both localities contain the same species of
animals (some species, however, were found in only one of the fissures). They are
probably of the same age. Besides mammals, the Przeworno fauna contained
numerous amphibians, reptiles, and some remains of birds (Mlynarski, 1978, 1984;
Szyndlar, 1984; Bochenski, 1987). The very scarce remains of small mammals (insecti-
vores and rodents) are so far unidentified. Primates, as in Opole, were represented by
PlioTiithecus antiquus (Kowalski and Zapfe, 1974). From rodents, only one species,
Cha1comys jaegeri, was described so far (Kubiak and Wolsan, 1986). Carnivores, only
a small part of which have been so far identified, contained Pseudaelurus
quadridentatus and P. larteti. The only proboscidean is Gomphotherium angustidens
(Kubiak, 1975). Among perissodactyls there were Aceratherium simorrense,
Brachypotherium brachypus, and Anchitherium aurelianense (Kubiak, 1981b); among
artiodactyls so far only Conohyus simorrensis, Taucanamo sansaniense, Euprox
furcatus, and Dorcatherium crassum have been identified (Kubiak, 1981a). The
composition of the Przeworno fauna suggests the presence of a differentiated
environment in the vicinity of the fossil locality, with both humid forests and savannas
(Kubiak, 198Z).
The absence of deposits in Poland containing terrestrial vertebrates representing

zones MN 8 - MN 13 is probably not accidental. During that time marine transgression
of the middle Miocene brought the deposition of sediments of Paratethys in the
Carpathian Foredeep. To the north, these deposits passed into the alluvial plain of the
Polish Lowland. The Poznaii. Formation, which was deposited there, exhibits numerous
marine influences (e.g., in the fauna of Foraminifera).
The erosion of a part of these sediments probably took place at the time of the
Messinian crisis. The sediments of Paratethys and clays of the Poznan Formation are
now separated by outcrops of older rocks of the Meta-Carpathian arch. The middle
Miocene layers were eroded as a result of Pliocene uplift and erosion in this area,
which started in the Messinian (G1azek and Szynkiewicz, 1987).
PUOCENE
The deposition of the next vertebrate fauna of Poland took place at the end of a
long arid interval in the entire Middle Europe, documented by many geomorphological
and sedimentological features in Poland. It is the fauna from Podlesice (PO) in the
Krak6w-Wieluii Upland. It was discovered by K. Kowalski in 1951 in calcite layers
deposited in a vertical cave. The fauna is dominated by bats; the remaining bones,
nearly exclusively those of small vertebrates, probably originated from owl pellets.
The fauna of insectivores of Podlesice is strikingly rich (Rzebik-Kowalska, 1971,

1975, 1976, 1981, 1988). Among them were no less than 1Z taxa of shrews; the moles
203
were also highly diversified (Skoczen, 1976, 1980). The Podlesice bats (Kowalski,
1956; WQ!oszyn, 1988) include genera living in Europe today. The presence of two
species of horseshoe bats (Rhinolophus) and of the genus Miniopterus points to a
climate of Mediterranean type.
Rodent fauna in Podlesice is rich and very peculiar (Agadjanian and Kowalski,
1978). There are there five species of sciurids (Black and Kowalski, 1974), and three
taxa of eomyids includin~ the genera Estramom~s, Keridomys, and Leptodontomys
(Fahlbusch, 1978). Podles1ce is tbe type Iocahty oKowaiSkia polonica arid K. magna
(Fahlbusch, 1969), later discovered over a great area of Europe and Asia. The
presence of Gerbillinae, represented by the genus Epimeriones, may suggest an arid
climate (Kowalski, 1974). Besides hypsodont cricetids of unknown affinities (like
Microtodon), typical arvicolids (e.g., Prosomys insuliferus) were already in Podlesice,
recorded for the first time in the area of Poland. Murids are scarce; glirids are quite
diversified and numerous (Kowalski, 1963). Among zapodids, there is SminthoziB:us
intermedius, first described from Kohfidisch (Kowalski, 1979). The only specificly
identified carnivore from Podlesice is Baranogale helbingi (Kowalski, 1959).
A similar faunal assemblage is known from layers 4-5 of the Maia Cave in the
Krak6w-Wielun Upland. The stratified sediment contained Quaternary vertebrates in
its upper layers, but the oldest fauna is evidently only slightly younger than
Podlesice. Remains of small vertebrates were found exclusively and their number was
limited. Most of the species of bats, insectivores, and rodents were also present in
Podlesice; of particular interest is the presence of Epimeriones (Sulimski et al.,
1979). Another locality of this age, so far not studied in detail, is Paiiska Gora.
Elements of fauna of the same age, i.e., the early Ruscinian (MN 14), were found
in some other localities in Poland with mixture of animals of different age, e.g., in
Zamkowa Dolna Cave and Zalesiaki.
The late Ruscinian (MN 15) is best represented in Poland by the rich fauna from
Wfi!ze 1 (WEI), situated at the northern end of the Krak6w-Wielun Upland. Sediments
of a vertical pit of W-;ze have yielded bones of large as well as small vertebrates. The
climate during the accumulation of these fossil remains was relatively warm, of
Mediterranean type indicated by a rich reptilian fauna, including turtles (Glazek et al.,
1973).
Among the insectivores, there was a surprisingly large number of shrews, about
15 species (Sulimski, 1959, 196Zb; Rzebik-Kowalska, 1971, 1975, 1976, 1981, 1988), as
well as hedgehogs, desmans, and moles. Bats (Kowalski, 196Z) do not significantly
differ from those present in the earlier fauna. Among the lagomorpha, the leporid
Hypolagus beremendensis appears at W-;ze for the first time in the Polish fossil
record; it is a characteristic element of all later Pliocene and early Quaternary faunal
assemblages (Sych, 1965, 1967a,b). Fauna of rodents was very rich; it included five
species of arboreal sciurids, differentiated glirids, and some murids. This is the unique
locality with a fossil representative of the family Seleviniidae, a family now popu-
lating only arid environments in Asia (Sulimski, 196Za). The presence of mole-rats
(Kowalski, 1960c) is another indicator of a rather arid climate. There are very few
typical cricetids (Pradel, 1988); on the contrary, aberrant forms of this family, such as
Trilophomys (Schaub and Kowalski, 1958) and Baranomys (Repenning, 1968) with
hypsodont teeth are abundant. In the number of individuals, arvicolids are dominant;
they were studied by Kowalski (1960b) and Sulimski (1964), but a modern revision is
badly needed. Only forms with rooted molars are present; one of them, Stachomys
trilobodon, has many characters in common with the recent Prometheomys
schaposchnikovi from the Caucasus. Other taxa of voles recorded from W~ze 1 are
Ungaromys, Germanomys weileri, Dolomys cf. milleri, and Propliomys hungaricus.
The genus Mimomys was represented only by primitive forms, mainly M. gracilis and
M. hassiacus. There was in the material a fossil porcupine, Hystrix primigenia
(Sulimski, 1960).
204
The fairly rich fauna of carnivores is not very useful for stratigraphic purposes,
as the majority of taxa described from WS!ze 1 were new species; e.g., Felis wenzensis,
Agriotherium intermedium, Martes wenzensis, Mustela plioerminea, M. pliocaenica
(Stach, 1951, 1954a,b, 1957, 1959, 1961). The taxonomic position of these carnivores
needs further study. Some additional data about the fauna of carnivores may be found
in publications by T. Czy:2ewska (1961, 1969, 1978, 1981).
W~ze 1 yielded scarce remains of a rhinoceros, Dicerorhinus megarhinus

(Czyzewska, 1958), and two species of deer, Procapreolus wenzensis and Cervus
warthae, were both originally described from this locality (Czyzewska, 1959, 1960,
1968, 198Z).
The same period is represented by many faunal assemblages discovered in the

Krakow-Wielun Upland (e.g., Mokra 1, Draby 1, Ewy Cave, Raciszyn 1). These assem-
blages were usually rather poor and not studied in detail.
The next period which left traces of its vertebrate fauna in Poland is the Early
Vil~anyian (MN 16). Two localities, both probably of the same age, in RE(bielice
Krolewskie, RK1 and RKZ, that belong in this interval are of prime importance. They
contain mainly small vertebrates and are fillings of karstic fissures in Jurassic lime-
stone, situated a few hundred meters from one another.
The red color of the sediments and the composition of the faunas (e.g., the
presence of numerous reptiles) point to a relatively warm, Mediterranean-type
climate.
Insectivores were less diverse than in older faunas, only nine species of shrews
were present. Bat remains were rather rare; included among them were two rhino-
lophids. Hypolagus beremendensis is the unique lagomorph recorded (Sych, 1980).
In the fauna of R~bielice Krolewskie rodents (Kowalski, 1960a) there were

eomyids (Estramomys cf. simplex), beavers (Trogontherium), murids, and glirids. A
representative of zapodids, Sminthozapus janossyi, also recorded from W~:!e 1, occurs
here for the last time (Kowalski, 1979). Arvicolids are numerically dominant; they
belong mainly to rooted forms, e.g., Stachomys is still present; the genera Borsodia,
Dolomys, Propliomys, and Villanyia are also there; Mimomys is represented by several
species including M. polonicus, a species described from this locality and discovered
later in many other faunas. Of particular interest is the presence of the earliest
arvicolids with permanently growing, rootless molars, e.g., Synaptomys europaeus
(Kowalski, 1977). This genus belongs to lemmings and is limited in the recent fauna to
the boreal zone of North America. The hypsodont cricetids Trilophomys and
Baranomys are also present. Only one carnivore, Baranogale helbingi, is known from
R~bielice Krolewskie 1. There were also scarce remains of deer, identified as
Croizetoceros ramosus and Eucladocerus sp. (Czyzewska, 197Z).
The same period is also represented by the locality W~~e Z (WEZ), the
mammalian fauna of which has not been studied in detail. According toT. Czyzewska
(pers. comm.), cervids are diversified and four species were represented in the
material, which was composed mainly of teeth. There were Cervus cusanus, C.
pardinensis, Arvemoceros ardei, and Croizetoceros ramosus. Also, locality Mokra Z
brought some rodent remains of similar age.
There are also two localities in the Krakow-Wieluii Upland with rich faunal
remains, but with a mixture of material from different age. One of these is Zamkowa
Dolna Cave, situated inside a medieval caste!. During excavation, the original order
of layers was destroyed. The fossils represent an interval from the early Pliocene to
the late Pleistocene, but the fauna of the Villanyian is dominant. An ochotonid,
Ochotona polonica, was described from this locality (Sych, 1980). Zalesiaki is another
locality with similar faunal elements.
205
Table z. The appearance of selected species of rodents in the Polish faunal assem-
blages from the limit of the Neogene and Quaternary. Locality abbre-
viations: KI3B = Kielniki 3B; KD = Kadzielnia; KA = Kamyk; Kl3A =
Kielniki 3A.
Localities
Species Kl3B KD KA Kl3A
Spermophilus polonicus (Gromov, 1965) X X

Allocricetus bursae Schaub, 1930 X X X X
Allocricetus ehiki Schaub, 1930 X X X X
Cricetus runtonensis Newton, 1909 X X X X
Clethrionomys kretzoii (Kowalski, 1958) (sp.) X X X X
Lemmus X X X X
Ungaromys nanus KorlJ!os, 193 Z X X X X
Pliomys episcopalis Mehely, 1914 X X X X
Borsodia hungarica (Kormos, 1938) X X
Villanyia exilis Kretzoi, 19 56 X X
Mimomys tornensis Janossy et Vander X X X X
Meulen, 197 5
Mimomys pitymyoides Janossy et Van X X X
der Meulen, 197 5
Mimomys pliocaenicus (F. Major, 1889) X X X
Mimomys cf. reidi Hinton, 1910 X X
Microtus (Allophaiomys) X X X
NEOGENE-QUATERNARY BOUNDARY
The age of faunal assemblages from Poland which could represent the latest part
of the Pliocene (MN 17, Late Villanyian) is difficult to evaluate. There are four faunal
assemblages approximating this age in Poland: Kielniki 3B (KI3B), Kadzielnia (KD),
Kamyk (KA), and Kielniki 3A (Kl3A). The elements of their mammalian fauna have
not been listed in table 1. Some elements of their rodent fauna are enumerated in
table z.
The faunal assemblages mentioned above represent such a short interval of time
that the differences among them depend more on migrations under the influence of
-changing climate than on evolution in situ. Allophaiomys, a primitive subgenus of
Microtus that is generally recognized as an index of the beginning of the Quaternary,
is absent in Kielniki 3B, rare in Kadzielnia, and is common and diversified in the
following two localities (Kamyk and Kielniki 3A).
In general, the interval represented by all these faunal assemblages was

distinctly colder than the preceding interval. However, this interval was probably
warmer than the present climate of central Poland. There were rather numerous
reptiles present, including the glass-snake Ophisaurus pannonicus, a genus now
restricted in distribution to the Mediterranean area.
The majority of species recorded during that interval are today associated with
the European forest biota. Individual elements of the new fauna appear successively,
one by one. The number of insectivores diminishes; there are only four species of
shrews in the rich fauna of Kadzielnia. Flying squirrels disappear and ground-squirrels
(Spermophilus polonicus) make their appearance. There appear, from the beginning of
this period, the cricetids, which will be typical also for the later periods of the
Quaternary, e.g., Allocricetus ehiki, A. bursae, and Cricetus runtonensis. The genus
Lemmus is represented by a single species evidently not yet limited to the Arctic
environments. The number of rooted voles diminishes. The evolutionary line of the
large species of Mimomys is represented by M. pliocaenicus; the populations from
206
Kamyk and Kielniki 3A are more derived than earlier ones (A. Nadachowski, pers.
comm.).
The drawing of the line dividing the Neogene and the Quaternary in Polish
mammalian successions must be rather arbitrary.
As could be seen, Polish localities of fossil mammals from the Neogene are
mainly connected with the fossil karst. This makes their correlation with general
stratigraphy rather difficult, and therefore they cannot contribute much to the
Neogene stratigraphy. On the other hand, many of them are rich in remains, particu-
larly those of fossil small mammals, and in some cases they permit a characterization
of populations from different stages of evolution.
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Rzebik-Kowalska, B., 1976. The Neogene and Pleistocene insectivores (Mammalia) of
Poland. m. Soricidae: Beremendia and Blarinoides. Acta Zoo1. Cracov., v. Z1,
P• 359-385.
Rzebik-Kowalska, B., 1981. The Pliocene and Pleistocene Insectivora (Mammalia) of
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Acta Zool. Cracov., v. 3Z (in press).
208
Schaub, S., 1925. Die hamsterartigen Nagetiere des Tertiars und ihre lebenden
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209
THE NEOGENE VP SITES OF CZECHOSLOVAKIA:
A CONTRIBUTION TO THE NEOGENE TERRESTRIC
BIOSTRATIGRAPHY OF EUROPE BASED ON RODENTS
Oldnch Fejfar
"" ~ y..... ...
Ustredm Ustav Geologiclty
Malostranske Namest19
Praha 1, Czechoslovakia
INTRODUCTION
The Tertiary VP sites of Czechoslovakia cover the time span since late Eocene
with gaps in the middle-late Oligocene and late Miocene (MP ZZ-MN Z; MN 7, MN 8,
and MN 10-13). The Neogene faWlal sequence includes occurrences of general impor-
tance: (1) the MN 3b faWla of the riparian gallery forest in the base of the brown coal
seam in the north Bohemian Tertiary basin, (Z) the superposition of the 4 VP assem-
blages in the profile of the Cheb Tertiary basin in the hanging sequence of the main
brown coal seam: Dolnice 1-3 (MN 4a-MN 4b) and Franzensbad (MN 5); the faunas
belong to the same type of environment, (3) the marine littoral-deltaic MN 4 faWla of
Orechov - correlated with Dolnice 3 -- underlying the Rzehakia (Oncophora) beds,
contributes to the correlation of the MN 4 faWlas with the Ottnangian stage of the
Paratethys, (4) the faWlal sequence of Dev{nska Nova Ves (Neudorf an der March) is
directly linked to the near coastal environment of the Badenian stage; the karstic
fillings (Spalte 1-3) with rich small and large mammalian remains are twice trans-
gressed: with sediments of two subsequence zones: (a) Lageniden zone and after a
short regression (b) Bulimina-Bolivina zone; the deltaic littoral sands of the latter
transgression (site Neudorf-Sandberg) produced large mammalian remains.
The intermediate position of the faWla of Franzensbad between Dolnice 1-3,

Orechov, and the fillings (Spalte 1-3) of Devinska No.va Ves is proved by the evolution
of several cricetid taxa: Neocometes, Anomalomys, Democricetodon, Mega-
cricetodon. This makes the (lower) Badenian age of the karstic fillings more probable
than the Karpatian one.
The Czech MN 4-5 Neogene VP faWlas contribute further to the knowledge of

extensive changes: extinction of Melissiodon, Ligerimys, and Pseudotheridomys, and a
subsequent immigration-replacement of Democricetodon, Megacricetodon, Eomyops,
Keramidomys, Neocometes, Anomalomys, and Eumyarion. To those changes, the
Proboscidean datum seems to be closely related (the first record of Gomphotherium in
Dolnice 3, and of Deinotherium in Franzensbad).
Karst fissures with the Vallesian MN 9 faWla in middle Bohemia at Suchomasty

fill a gap in the middle Miocene and Pliocene record; the assemblage contain a
mixture of typical Miocene genera together with several new immigrants, ancestors of
the modern European faWla.
The late Pliocene assemblages (Ivanovce, Hajnacka, Vcelare, Koli~any, Plesivec,

Ctfueves) contribute together with HWlgarian, Polish, and Austrian sites to the
N
~
N
Fig. 1. Review of the biostratigraphical position of Czechoslovakian Tertiary VP
sites, Part A: West and North Bohemian localities; Part B: Middle and
South Bohemian, Moravian, and Slovakian localities. Map of
Czechoslovakia showing the position of localities in Bohemia, Moravia,
and Slovakia. Bohemia: 1 = Valec, Dverce, Pet.aii 1,Z; Z = Chomutov
region - drillings Ah, Co, Kr, etc., brown coal J»it Merkur-north; 3 =
Cheb basin - Dolnice 1-3, Frantiskovy La~e (Franzensbad); 4 =
Lukavice; 5 = Tucborice - site 1: limestone quarry, site Z: hot spring
travertines (Suess and Reuss); 6 =Czech karst: Ko~eprusy {fillings C 718,
JK 1-4), Suchomasty, Chlum; 7 = Strakonice, L{S9v• Moravia: 8 =
oi"echov; 9 = Milrulov; 10 = Hovorany, Mqtenice. Slovakia: 11 = Devfnska
Nov~ Ves (site 1: Neudorf-8palte 1-3, Bonanza; site Z: Neudorf-Sandberg
sand pit; site 3: Neudorf-clay pit of the brick fabrique); 1Z = Ivanovce;
13 = Ko$any; 14 = Haj~lltka; 15 = Gombasek, Plesivec; 16 = VceJare.
·~
STAGES STAGES
PLANCTON .......,
zones MAM\tAL
"';;! ~~
NN;.:; M%,
CENTRAL
BOHEMIA MORAVIA SLOVAKIA
MEDITERRANEAN AGES
"t
::
:1: PARATETHYS
PLEISTOCENE PLEISTIICENE ~
N22 BIHARIAN • ~~!~r~st c7te, JK • Str6nsk6 skllto
Ho!s.te·n I Gombosek
PIACENCIAN N21 m MN17 • Ctinivu - JovoriCko ~1~~ny, ceo\Ore
~
VILLANYIUM
ROMANIAN I - ~ MN1& • Hornomoravsk)' uvol
• HajnO:Ho t2
~ ZAJICLEAN N~
20 ~ ~15b RUSCINIUM
• I vanovce \2
DACIAN
5
MESSINIAN
hni NN12 MN14
....._.,_
MN12
PONT IAN
N1'1 NN11
- TUROLIAN
MN11
~~IAN -
N16 NN10
MN 10 IMuUnice
10 Aa.NNONIAN VALLES IAN
MN~ • Suchomos ly • Hovorony (Kyjovseam
~ NN9
N14
Nii"f MN 8
SERRAVALLIAN SARMATIAN ~13 NN?
MN 7 ASTARACIAN
N12
~ • Mikulov k
~ ~~~n::rtl.. ova es
~::f
~ 1-- ~·
15 MN 6 • Devmskl ov6 Yes
BADENIAN NNS S alt 1-3
LANGHIAN
::0:
KARPATIAN ~ e-----,
5
I
• ~~~0n~~~i~:ad
I
NN4 Dolnice 3 • Ofechov
N?
OTTNANGIAN - ____.:: MN4 Dol nice 2
BURDIGALIAN f;s NN3 ORLEANIAN

Dol nice 1
~-"-" 1-- b
.
20
] EGGENBIJRGIAN
N 5 NN2
MN3
Tuchofloo
Merkur-Nor.th
Drillings: Ah,Co,Kr
AQUITANIAN N 4 NN1 MN 2
AGENIAN
.
Q
25 p 22 NP25 :~,~
:3 CHATTIAN
EGERIAN - - MP29
MP28
MP27
p 21
""'26
NP24 jMP25
30 ~ jMP24 SU EVIAN
~j RUPELIAN -
~
1--
MP23
~ NP:!:
- f;;;, MP22
KISCELIAN p 19
35
] LATOORFIAN
p 18 I--
MP21
I oitan, vatiC
overtf', Lukovic.e
- NP21 ~P20
I--
t.n, ~~
MP19 HEAOONIAN
P17
PRIABONIAN NP~
~ '•' 1...18
Fig. z. Biochronologic distribution of the Czechoslovakia Neogene VP sites.
213
sequence of the MN 14-MN 17 levels, and to the recognition of the Pliocene-
Pleistocene boundary.
The Neogene VP sites of Czechoslovakia (figures 1 and 2.) occur (1) in the
erosional relics of the Bohemian massive, (2.) in the region of Karpatian fore-deep on
the eastern slopes of the Bohemian massive, (3) in the northern part of the Vienna
basin in southern Slovakia; (4) numerous sites of late Pliocene and Pleistocene age are
scattered in different karst regions.
REVIEW OF THE SITES
Sites of Bohemia
The Bohemian massive is an old geological nucleus of middle Europe, uplifted

and intensively eroded since the last marine transgression of the late Cretaceous.
During the Paleogene, the whole area was subsequently leveled in a peneplain; at the
same time, in the north and northwest part, a rift structure of southwest-northeast
direction produced intensive alkalic volcanism. Several basins and depressions along
the rift were filled with debris, volcanic agglomerates, and, locally, with brown coal
seams. The deposits contain many fossil sites, including superposed paleobotanical
sites younger than late Eocene, and several levels of mammalian faunas.
Valec, Dverce, Detaii, aud Lukavice: (early Kiscelian, early Oligocene), early
Suevian local faunas and assemblages of the MP 2.1 zone. All sites are in context of
the initial volcanic activity of the rift, in subaeric ashes (e.g., Detan on the base of
the stratovolcano of Doupov Hills, radiometric data: 37.7 :1: 1.5 Ma), or in redeposited
limnic fine tuffites and seam layers with floras and molluscs (Fejfar, 1987).
Region of Chomutov aud Most, north Bohemia: (Eggenbut;gian, ~arly Miocene),

early Orleanian assemblages of the MN 3 zone, in drillings .(Ctyroky et al., 1964;
Fejfar, 1974) and in the same level, exposed in the nearby open coal pits (e.g., Merkur-
north); all in the seam deposits of the main brown coal seam - lignitic calcareous
pelites, marls, with abundant limnic and terrestric molluscs. The same level of differ-
ent lithology and paleoecology is exposed in the southern margin of the basin, as fault-
bounded blocks of bedded limnic limestones (Tuchorice - limestone quarry with
molluscs; Klika, 1891; Wenz, 1917), or erosional relics of hot spring travertines with
abundant mammalian remains (but without molluscan fauna; Tuchorice - site of Suess
and Reuss; Fejfar, 1974, 1985). .
Cheb basin: Dolnice 1-3, Franzensbad (Ottnangian, Karpatian, late early

Miocene), late Orleanian assemblages of the MN 4a,b and MN 5 zones. In a hanging
sequence of the main seam, in calcareous marls and silts (Fejfar, 1974).
South Bohemian Tertiary basin: two assemblages: Strakonice (Karpatian, late

early Miocene) of the MN 5 zone, related to Franzensbad; drillings at L1sov
(Sarmatian, middle Miocene) of the zone MN 8 related to the sites of the Upper
Bavarian molasse (Giggenhausen) (Fejfar, 1974).
Suchomaaty: middle Bohemian karst in Paleozoic limestones (early Pannonian,

late Miocene), Vallesian rich small mammalian assemblages of the MN 9 (or MN 10)
zone; in stratified fine-coarse quartz sands and clayey sands in karst fillings (? relics
of subterranean karstic streams).
Ctfneves: middle Bohemia (latest Pliocene), Villanyian,.. ,

fauna of the MN
,#,.,
17
zones; stratified sequence of slope debris of a basalt cone R1p (Fejfar and Horacek,
1983).
Kon;prusy, Chlum: (early Pleistocene), Biharian faunas in stratified cave-

fissure deposits in the Czech karst (Fejfar, 1961, 1976; Fejfar and Horacek, 1983).
Prezletice, early Pleistocene, late Biharian site, northeast of Praha; in limnic marls
214
I~
w MAMMAL
>: STAGES STAGES Distribution of rodent taxa
::I CENTRAL AGES
MEDITERRANEAN
:f PARATETHYS
~
IN23~ -~ii Ill
PLEISTOCENE PLEISTOCENE c c >-111
tN22JNN 19 BIHAR IAN :0 ~-~ ~~
LU PIACENClAN
~MN17
N 21 NN b VILLA NY IAN ·-
~;
~-r--~
H .
~g ~~
dH~
O.w~ ~l~
IHII
I
16
§ ROMANIAN NN 15 MN16a ] w ~ ~ ~-~i H?rl ' 1
11 111
3 19 ""N"Nt4 MN 15 b ·a g_ o
~ t"a. e ~I 1 1
a.. ZANCLEAN N2-0TI, a RUSCINIAN .; ~~~g:i ~~:.":.
>. >.-, > :1:- 10
~-ow1 .!:!~'IIIII
S I DACIAN h:;-;;; NN 12 MN 14 1'1 ii~ ~. ·•EEE-aE
a
1111 ~ ee ~.
1 N 18 111
.;~-C ~.; ~ ~-~~~(;&
~ MESSINiAN fMM..1L :; : -~ 4-t---r-~~e-~ ~ 111~ ~~~~-;~-~~i-~~~m5
N 17 MN 12 ~~ [ g:5~ 111
~-5~n.u ~ ~ ~~~]i_~ g~5~~~
PONTIAN NN11 1--- TUROLIAN •.: - ,·.:·.: ~ ~.c 1111 ·~·~;;• e!!a::o.e
~~ ~ ii ~~ ~ ~~~
;;-~ &~~ uJ: ~Vlmeg~
c u
. .,...___
t---
MN11 ~:.:;:
:1:<!1 c(
:~"'
~~ a
~ ~-~a:
o.:l:
,g..c:
a:
n.~
~I TORTONIAN
N 16 NN10 MN 10 111 I ~~ 111 ~ ~~I ::_
I I I ~I i I E
10 PANNONIAN VALLESIAN ;: • '-''II g
N15 MN 9 I E' r11 ~--!1
I - - NN 9 ; ~
N 14 " :; :; '
I ~ MN 8 ~ ~ !:'-!L·~.E 0:~
NN 8 I---~ r- -~ =e _, ~ -~
SERRAVALLIAN SARMATIAN N 13 NN 7 MN 7 •~ • ·: 1 ;;;
~ 1-- ASTARACIAN 1---~ r-- f; 1 w
'!'! N 12 NN 6 c 1'.• o !;
~~ t-;::r;T J-- MN 6 j Ul fg. ~ g~
15 -IUJ BAOENIAN ~ -~~ -~~ ~ "' z
0 LANGHIAN ~ NNS >'"-• i:l) .~ !. IJ
o - MNs :1: ~e o.. r• _, !'" 1·
:::;: KARPATIAN IN- 8- 'NN4 r---b ~~~~·:1e ::"-1
OTTNANGIAN N7 r-- MN 4 a ~ ~ull I II I
BURDIGALIAN t;s NN3 ~ ORLEANIAN ]
.20 ~I
a EGGENBURGIAN I N 5 NN 21MN 3 I
w
a
bl I II
AQUITANIAN I II
IN 41NN1 rN 2 al AGENIAN
1--+------1 EGERIAN ( p. I
25 CHATTIAN (p. I
p 221 NP~~/~P~ SUEVIAN ( p) tllllll I
Fig. 3. Chronologie distribution of important taxa of Rodentia in the
N middle European Neogene, based on Czechoslovakian record
....
U1 1957-1987.
with mollusc!b large and small mammals (Fejfar, 1969) and early acheulian Paleolithic
implements (Sibrava et al., 1979).
Sites in the Karpatian Fore-Deep
During the early and late Miocene, the Karpatian fore-deep has been repeatly
transgressed by the Paratethys sea from the south. Thus, the paralic conditions on the
southeast slopes of the Bohemian massive provide possibility to find mammals in the
near coast oligohaline deposits. Later during late Miocene (Pannonian and Pontian),
the south Moravian region charged into isolated brackish-freshwater basins surrounded
b'l gradually flooded swamps and forests producing locally lignite seams (Hovorany,
Sardice).
Oiechov, south of Brno: (late Ottnangian, late early Miocene), marine-

estuarine, fine- to medium-grained, calcareous, glauconitic sandstone with layers of
cross-bedded fine ferruginous sands; on the base coarse gravel sandstone over crystal-
line metamorphic rocks. Thickness of the outcrop is 3-4 m. In the fauna are rarely
ribs of sirenians and shark teeth. In the close vicinity, in drillings and occasional out-
crops, this type of ~deposit is overlain by Rzehakia (Oncophora) beds of the late
Ottnangian stage (Ctyroky, 1968, 197Z, 1973, 1987; Cicha et al., 197Z). The
mammalian fauna is comparable with the upper level of Dolnice (Dolnice 3) of MN 4b
zone.
Mikulov (Nikolsburg}, south of Brno: (late Badenian, Sarmatian, middle

Miocene), fine- to medium-grained sands on the top of the marine late Badenian
deposits produced large mammals only of the zones MN 6 and MN 7.
Mutemce, Hovorany, southeast of Brno: (Pannonian, late Miocene), two faunal

levels in the Kyjov lignite seam (Pannonian B) in Hovorany, and the hanging sandy
clays at Mutenice (Pannonian D; small mammals).
Karstic fissure fillings JavPi~Ao: (late Pliocene, MN 17), Stranska skala,

Holstejn (early Pleistocene, Biharian); limnic sandy marls in Hornomoravsky uval (late
Pliocene, MN 16b). Rich small mammalian assemblages with typical arvicolid taxa
(Musil, 1966; Fejfar and Horacek, 1983).
Sites in Slovakia
Devlnsu Nova Ves (Neudorf an der March), figure 4: The near coastal region of
the Paratethys sea during Badenian in the northern part of Vienna basin northeast of
Bratislava. Several importrant localities of invertebrates plus land and marine verte-
brates (Zapfe, 1949; Papp et al., 1978), all intercalated with two subsequently trans-
gressive zones of marine early and late Badenian (Lagenid zone; Bulimina-Bolivina
zone): Karstic sites: Spalte 1-3 and Bonanza (early Badenian-Moravian, early middle
Miocene); tectonically produced fissure fillings combined with the ancient cave system
(Zapfe, 1949; Fejfar, 1974). Site Bonanza, partly with bedded medium-grained
phosphatized sandstones containing marine fishes and mammals, probably represent
deposits of a submarine cave (Holec et al., 1987); all karstic filling belong to the same
early MN 6 level; younger age than lower Badenian-Moravian is excluded by the
marine transgressions (covering also the karstic Jurassic limestones and partly filling
their upper parts), an older Karpatian age is excluded by the evolutionary level of
cricetids (figures 8-11). Neudorf-Sandberg (late Badenian-Kosovian, middle Miocene),
sequence of transgressive sands, sandstones, lenses ,.of cross-bedded estuarine deposits
(Papp et al., 1978) with marine molluscan fauna. (Svagrovsky, 1974), otoliths (Holec,
1978), and marine and land mammals (Thenius, 195Z). The whole sequence in a former
large sand pit (now a natural reservation) represents a littoral, near coast facies of the
Bulimina-Bolivina zone of the late Badenian; the mammalian level belongs to the late
MN 6 zone. Neudorf-clay pit of the brick fabrique; marine clays (Badener Tegel) with
invertebrates Amusium cristatum badense and Discoaster exili~, plus land flora in
indurated claystones; nannoplankton of the NM 6 zone (Lehotayova, 1977). Deposits of
the clay pit are in a tectonically faulted block, and represent the stratigraphically
youngest Badenian level in the area.
216
Loca liti es of Sandberg
Devinsk6 Novo Ves
( Neudorf o.d. March) ----------l
:-:-:-:-;-·;-,:=-_-:_
2km to the E
Old sand p i t
----------r45m I
r
60 m
Early Sar matian level
___L ______
2 ]{§.~~{}?~')
3 E
4 IT!-
5 ~
Om
6 m
Fig. 4. Geological sections of localities in Devfnska Nova Ves: 1 =limestone

quarry with several karst fillings in Jurassic limestone (Neudorf-Spalte
1-3, early Badenian, lower Astaracian assemblages of the MN 6 zone);
2. = sand pit Neudorf-Sandberg (sequence of transgressive gravels, sands,
and sandstones on Jurassic limestone; late Badenian, Kosovian assem-
blages of marine invertebrates and both marine and terrestrial large
mammals of the late MN 6 zone); 3 = clay pit of the brick fabrique
("Badener Tegel" clays of the latest Badenian, Kosovian in tectonically
foundered block). Explanation of the deposits: 1 =bands of sandstones
with Lithothamnium, 2. = polymikt psephites, medium-coarse grained,
3 = yellow-grayish fine to medium grained sands, cross bedded, with
vertebrates, 4 = gray clays of "Badener Tegel," 5 = coarse sands and basal
gravels, 6 = Jurassic-Cretaceous limestones with holes of the boring
molluscs (Lithophaga), with tectonically-altered karstic fissures.
Late Pliocene localities: Ivanovce, Hajpilcka, Vcelare, Kolmany, Plesivec,

cover the time span of MN 15b-MN 17 (Fejfar, 1961, 1976; Fejfar and Heinrich,
1983). Ivanovce, system of karstic fissures in Triassic limestone, two units of fillings
(older horizontal A, younger vertical B) both with late Ruscinian assemblages of large
and small mammals (Fejfar and Heinrich, 1985). Hajnacka, a sequence of lacustrine
sands, agglomerates (tuffs and tuffites), probably a filling of a volcanic maar; the
whole site in direct relation to a basaltic eruption (Fejfar, 1964; Fejfar and Heinrich,
1985), large and small mammalian fauna of MN 16a (early Villanyian). Flora in
tuffites overlying the mammalian level. Ycelare, Kolfiiany, Plesivec, all karstic sites
with the latest Pliocene , Villanyian, small mammalian asse mblages of MN 17 zone . In
a limestone quarry Vcelare, southern Slovakia, a complex system of fillings (some of
them stratified) Vcelare 1-7 represent a sequence from early Arondellian MN 16a to
early Bihatian faunas (Fejfar and Horacek, 1983; Horacek, 1985). Gombasek in south
Slovakia, Zirany in middle Slovakia, produced rich small mammalian assemblages
(Fejfar and Heinrich, 1983); Gombasek locality is of historical importance for the
2 17
early Pleistocene mammalian biostratigraphy because of the first record of the joint
occurrence of small and large mammalian taxa (Kretzoi, 1938, 1956; Fejfar and
Horacek, 1983).
BIOSTRATIGRAPMCAL INTERPRETATION
The Czechoslovak Neogene mammalian sequence documents the following data

of biostratigraphical importance (Cicha et al., 197Z; Fejfar and Heinrich, 1981, 1983,
1987). Based on the occurrence of mammalian assemblages in stratigraphical succes-
sion, some of them in marine or paralic sediments. This provides - along with evolu-
tionary changes of rodents - valuable clues to the age of parts of the mammalian
sequence.
Lower Miocene, early Orleauiau, MN 3 assemblages. Overlain or on the base of

the main brown coal seam (drillings around Chomutov, open coal pits); freshwater
limestones, hot spring deposits (travertines). Typical localities: coal pit Merkur-
North, Tuchorice.
Characteristic rodent taxa of the sedimentary rocks (pelitic calcareous sandy

marls with molluscs) of the seam: frequent Plesispermophilus descendens, Ligerimys
lophidens, Pseudotheridomys parvulus, Myoglis antecedens, Melissiodon dominans; rare
Blackia, Glirudinus, Miodyromys, Peridyromys; very rare Ameniscomys,
Heteromyoxus, Eomys, Miopetaurista, Microdyromys, Bransatoglis, Ptychoprolagus,
and Amphilagus. Characteristic taxa of molluscs: Nystia rubeschi, Gyraulus
trochiformis aplanatus, Planorbarius, Milax crassitesta, Sphaerium, Carychium,
Carychiopsis, Pyramidula, and Cepaea. Paleoecology: riparian mixed mesophytic
gallery forest with shrubs in understories (higher proportion of evergreen trees), along
gradually flooded swamps of the coal-forming vegetation with Glyptostrobus and
Taxodium.
Characteristic taxa of rodents of the contemporaneous hot spring travertines:

frequent Heteroxerus costatus, Spermophilinus bredai, Ligerimys lophidens; rare
Pseudotheridomys parvulus, Ptychoprolagus, Glirudinus, and Peridyromys. Forest
along shallow water ponds-lakes with palms (Phoeni~ Livingstona), Acer, Ulmus,
Mastixiaceae(?), ~ Zelkova, Toddalia, Smilax, and~·
Differences in the synchronous mammalian assemblages indicate differences in

the biotope. We may conclude that the absence of Plesispermophilus, Myoglis, and
Melissiodon in the travertines (with the travertine dominant Heteroxerus costatus
missing in turn in the coal marls) indicates the absence of fruit-bearing vegetation in
Tuchofice. Eomyids occur equally in both faunas and are of the same evolutionary
level. Moreover, the assemblage of Tuchorice (predominant large carnivores in the
macrofauna as a cause of the trapping effect of hot springs) may represent a
thanatocenosis.
Both assemblages have their closest affinities with the rich fauna of the karstic
fissure at Wintershof-West (Dehm, 1950). The faunas of the MN 3 zone, with absence
of the bunodont cricetid genera cover the time span of Eggenburgian.
Late lower Miocene, late Orleauiau, MN 4a, b, aDd MN 5 aaaemb:lages. Above

(in the roof of) the main brown coal seam in the Cheb basin. The vertebrate sites are
in a sequence (superposition) in one profile, in environmentally similar but climatically
changing (gradual deterioration, decrease of humidity) conditions:
Franz ensbad MN 5
Dolnice 3 MN4b
Dolnice Z
Dolnice 1 MN4a
218
(I@Sif. ···~~~
.;;
'" .---.. .==-1""".~33516~~ ~~~ ~'
~~~\l:i)~~~ .. 14 15 16 17 :;,?-'
~5 6 7 8 1333"' ~1333136 1
Fig. 5. 1 ~ I' " -
Meaaur~ts and represen- ° mm '

tative specimens of eomyid gen- ~ ~
era Ligerimys (Stehlin and
Schaub, 1951) and
WI
8 1~
fi' ~ ~
1a
~---
•
9
Pseudotheridomys (Schlosser, ~ W
'-'t p~
1926); M/1-2 (1-9), P4/ (10- u ..,
m1-2
20) with anterior of tooth g•
0
0 0
di~ected toward top of figure 0
\.3
in figures 5-15. 1, 2, 5, 10 .. .,0
\.2 11 a
Ligerimys lophidens (Dehm, 0
1950) from Tuchorice; 3, 4, 11, •• ...

12"' Pseudotheridomys ,parvulus 0 : ~ O/ '·'t
0 0
\.1 .. !' : ..
=· .·~ ooo
(Schlosser, 1884) fram the • 0
b. ..,og
base of the main brown coal 0 • 0 •
seam at Chomutov (3, 11) and .,...-!'-"" •

oA- 7333
0(1-- 7335
(.: '·
Dolnice (1, 4, 12); 5-8, 13-19 lO ~ ... - 7343
1.\l \. ": ..
• •
1. .
• Ligerimys florancei B• - 7363.7355.7382
(Stehlin and Schaub, 1951) •

•
\.0
from Dolnice 1 (5, 6, 13, 14), L L
1.\ \.2 \.3 \.5 mm- LO \.1 1.2 L3 ... mm
...., Dolnice 2 (7, 15, 16), Dolnice "
....
CQ
3 (8, 17-19)_, and Ofechov (9, 20) •
N
N
0 ' "'
-'
'
l 127 \-/,..,~/
~
1 2
8
Fig. 6.
Measurements and represen

tative record of eomyid
genera Keramidomys
(Hartenberger, 1966) and
Eomyops (Engesser, 1979);
PI 4 ( 2, 11 , 12) , PI 4-MI 3 ( 5,
6), P41-M/3 (13), MI1-MI2 10
(1, 3, 4, 7-10), M11-M21
~-
(14-26). 1, 2,14: ~~ 0 1 2 13 -
Keramidomys cf. thaleri ~ ~-.;/;
c=/·
~ mm
(Hugueney and Mein, 1968) ~
/
from Franz ens bad ( 1, 2)

and Strakonice (2); WI
Keramidomys carpathicus WI m 1-2 1-2
m-
(Schaub and Zapfe, 1953) 09
09t
from Dev!nska Nova Ves, / .~ ..
Spalte 1 (3-10, 12, 15-25); 12 -
11, 12, 26: Eomyops aff. 26
0.8~ I~~~-~·-· ··+·MT -;-
catalaunicus (Hartenberger, "l( ... X .
. ... . . :. •. ~
1967) from Strakonice (11) '\ • Keramidomys carpathicus . . . . ..
• Keramidomys cf tha/en· • - 7330
and Franzensbad (12, 26).
• Leptodontomys sp ... - 733051
07 07 t ~
L L
07 08 0.9mm 06 07 QB 0.9 mm
11
5 8
0 1 2 3
mm ~
12 13 14
2,3' w w
mr ~31 m~
2,2 ..
2,2
2,1
2,1
2fJ
2,0
1,9
c- Dolnice 2 1,
1,8 •- Dolnice 1
o- Orechov 1,8
I
1,7 •
L L
2:Z 2.3 2/. 2.5 2,6 2;/ 2,8 mm 2,7 2,8 ~9 3,0 3,1 3,2 3,3 3,4 mm
N Fig. 7. Measurements and representative record of Melissiodon dominans (Dehm, 1950); M/1 (1-8), M1/ (9-14) from the base of the
N
main brown coal seam at Chomutov (1), Dolnice 1 (Z-5, 9, 10), Dolnice Z (6, 11), Dolnice 3 (8, 13, 14), and Orechov (6, 7, 1Z).
In comparison with the nearby MN 3 faunas (at the base of the main coal seam)
distinct changes exist: new presence of Democricetodon, Eumyarion, Anomalomys,
and Neocometes in the levels of Dolnice 1 and 2.; new presence of Megacricetodon and
fragments of mastodon molars (cf. Gomphotherium) in Dolnice 3; after Dolnice 1 there
are also changes in the eomyids: Ligerimys florancei abruptly replaces Ligerimys
lophidens; the primitive line of Pseudotheridomys parvulus continues until Dolnice 3,
where it occurs together with Melissiodon and Myoglis. The record of Anomalomys
minor is very rare; only one molar in Dolnice 1.
Characteristic taxa of rodents in Dolnice 1-3 (the number of molars is given in

the order: Dolnice 1, Z, 3 in parenthesis): Pseudotheridomys parvulus (in Fejfar,
1974: Ligerimys antiquus) (2.4, 2., 3); Ligerimys florancei (118, 80, 83); Eumyarion
weinfurteri (18, 6, 42.); Democricetodon franconicus (ZZ, 36, 75); Neocometes similis
(4, 4, Z), Megacricetodon (Dolnice 3 only: 4); Melissiodon dominans (ZOO, 10, 67),
Anomalomys (in Dolnice 1 only: 1). The glirids: Glirudinus, Microdyromys, Glirulus,
Glis, Bransatoglis, Pseudodryomys, Miodyromys, Peridyromys, Myoglis.
In the Karpatian fore-deep, in the locality Orechov, below the Oncophora

(Rzehakia) beds of the late Ottnangian, the mammalian assemblages have close affini-
ties to the level of Dolnice 3; also present are: Megacricetodon, Democricetodon,
Eumyarion, Ligerimys florancei, and Melissiodon (all with the same species), but
Neocometes, Anomalomys, and Pseudotheridomys parvulus are missing.
Another evidence of the close relation to the Oncophora (Rzehakia) beds is given
by localities in the Bavarian molasse: Rembach, Forsthart, and Rauscherod (Ziegler
and Fahlbusch, 1986); the assemblage, comparable to Dolnice 1-3 and drechov, are in
limnic freshwater beds (limnische S\itlwasserschichten) above the upper marine molasse
and Oncophora beds. The evidence from Bavaria support the extension of the MN 4
time span through Ottnangian and the lower part of Karpatian.
The youngest level of the mammalian sequence in the Cheb basin, the fauna of
Franzensbad, provides further distinct change; two taxa of eomyids of Dolnice -
Ligerimr florancei and Pseudotheridomys parvulus - are replaced by Keramidomys
cf. tha eri and Eomyops sp. Neocometes, Democricetodon, Eumyarion, Mega-
cricetodon, and Anomalomys show further gradual evolution; also, the last archaic
genus Melissiodon disappears. The changes between Dolnice 3 and Franzensbad levels
are further proof of the climatic deterioration (decline of temperature and humidity).
Similar assemblage of the Franzensbad fauna comes from a relict in the region
of the south Bohemian Tertiary basin; the site Strakonice (figure 1b). In this fauna is
the only record of Cricetodon meini, Eumyarion bifidus, and the oldest find of
Lartetomys.
The faunal level of Franzensbad provides multiple evidence for the interpreta-
tion of the subsequent zone MN 6: (1) it shows distinct changes toward the preceding
assemblage of Dolnice 3 (in the same profile); and (2.) it poses an evolutionary inter-
mediate position toward the level of the zone MN 6, providing evidence - together
with the site Langenmoosen in the upper Bavarian molasse - for the zone MN 5, and
for the early Miocene stage of Paratethys, the Karpatian (Cicha et al., 1972.), and
probably the beginning of Badenian.
Characteristic taxa of rodents in Franzensbad (and Strakonice) (number of

molars given in parentheses) are Keramidomys thaleri (7), Eomyops (1), Demo-
cricetodon gracilis (1 72.), Megacricetodon bavaricus (108), Neocometes similis (5),
Anomalomys minor (1), Eumyarion weinfurteri (19), Eumyarion bifidus (1) (in
Strakonice only), Cricetodon meini (2.) (Strakonice only), Lartetomys cf. zapfei (1)
(Strakonice only).
Early middle Miocene, early Astaracian, MN 6 assemblages of Badenian. The

sites of Devi'nska Nova Ves (Neudorf an der March) (figure 4). Here, during Badenian,
two subsequent marine transgressions of the Lagenid-zone (middle Badenian) and of
222
8
0 1 2
lw mm
"
" \ - .
L
mm
a- Oolnlc• 1-3, Fronnnsbad
o·.: b - Sanson
c - Nt-udorf- Spalt• 1,2
L
umm ,.
" .. ..
Fig. 8. Measurements and representative record of the M/1 (1-9) and M1/ (10-17) of the genus
Eumyarion (Thaler, 1966). 1-6, 10-13, 17: Eumyarion weinfurteri (Schaub and Zapfe, 1953)
from Dolnice 1 (10), Dolnice 3 (1, Z, 11), Oiechov (3, 4, 12), Franzensbad (6), Strakonice (3),
N and Devfnska Nova Ves, Spalte 1 (17); 5, 14: Eumyarion bifidus (Fahlbusch, 1970) from
N
w Strakonice; 9: Eumyarion leemani (Hartenberger, 1965) from Mutenice.
N
N
~
'~'" ~""" _.. """" ~""'~'~
/: . :...,_ A
~ ~'"'
~·
~I~
~', /'(" A- r-'L"
Fig. 9. 15 16 17 18
1 3
~ - ~
Measurements
tive records and representa-
of the M/1 of ~,,...
"' ~'" ' I ' ~~ ~o~ ~~-
Democricetodon (Fahlbusch, ~~
1964) and Kowalakia (Fahlbuach, /~~ ~ ~ ,--
1969). 1-6: Democricetodon ~ ~ , -~
franconicus (Fahlbusch, 1966) 6 1 = a 9
from Dolnice 2 (2), Dolnice 3

(1, 3, 4), and Orechov (5, 6); 7-
12: Democricetodon gracilis
,..
(Fahlbusch, 1964) from
Franzensbad (7-10) and
Langenmoosen (11, 12); 13-14: 11
- 12 13 14
Democricetodon mutiluis
(Fahlbusch, 1964) from
Langenmoosen; 15-20:
I w
mT
Democricetodon vindobonensis
(Schaub and Zapfe, 1953) from
Dev!nsk~ Nov~ Ves, Spalte 2; 21 ,. I . ErbrtsJNJIM
2 • Lo Rom~u
Democricetodon minor (Lartet, J - Fro r'!ZM$bod •- 7~P- Spolf•1
1851) from Sansan; 22, 23: ' • Lattgt"NP'IOO.S:M Iii - 7YfWI 110n Dtmgcriutpdpo
• - Dolf'fk• 3 ~(Sdl llopf•J
Democricetodon gaillardi a- DtJII'Iiu 1
• •- ()drui;e P
(Schaub, 1925) from Suchomasty; :1( - O!Kh~~
24, 25: Kowalskia cf. L

fahlbuschi (Bachmeyer and L mm
'" mm
... "
Wilson, 1970) from Suchomasty;
26: Kowalskia intermedia
(Fejfar, 1970) from Ivanovce A.
~
Fig. 10.
9 10
Measurements and representa-
tive record of Ml/ of
I i
Democricetodon (Fahlbusch,
1964) and Kowalskia (Fahlbusch,
1969). 1-4: Democricetodon
franconicus (Fahlbusch, 1966)
from Dolnice 1 (4), Dolnice 2
(2, 3), and Dolnice 3 (1);
11-13: Democricetodon gracilis 0 , 2
w
(Fahlbusch, 1964) from mm
Franzensbad (11, 12) and ·~
Langenmoosen (13); 14: ml
Democricetodon mutilus w
(Fahlbusch, 1964) from ml "
Langenmoosen; 5-8: L
Democricetodon vindobonensis ..,mm
(Schaub and Zapfe, 1953) from

Dev!nska Nova Ves, Spalte 2; 9:
Democricetodon minor (Lartet,
1851) from Sansan; 10, 15, 16:
Democricetodon gaillardi
(Schaub, 1925) from Suchomasty; ..,___b
17, 18: Kowalskia cf.
fahlbuschi (Bachmayer and L
Wilson, 1970) from Suchomasty; mm
19: Kowalskia intermedia " "
...., (Fejfar, 1970) from Ivanovce A•
....,
(11
N
N
0)
~- ~,~-~·
3 4
~ 'f
Fig. 11
Measurements and represen- ~ ~
tative record of M/1 (1-11)
I " "' ~
I
and M1/ (12-19) of the ge- ~ =- ~
nus Megacricetodon
(Fahlbusch, 1964). 1-8, s ~
12-18: Megacricetodon 9 10
8
bavaricus (Fahlbusch, '~
1964) from Dolnice 3 (1, 1 2
0
12, 13), Orechov (2-4), mm
Franzensbad (5, 6, 15, 16), tA w
Strakonice (14), - and ~ W
Langenmoosen (7, 8, 17 ,18); m!
9, 19: Megacricetodon '·'
minor (Lartet, 1951) from i m1 '~
Dev!nska Nova Ves, Spalte "f
u
2; 10: Megacricetodon aff.
debruijni (Freudenthal,
1968) from Mut~nice; 11:
1.0
Megacricetodon· similis ~·
(Fahlbusch, 1964) from D - 73-lJ
6 - 1337
I / ·~~ I - LttRomi~
L!sov. 2 aValdthtDrO$ 111 B
.. - 731.7 I 3 •Vitu.III'-Collo,.,...s
/ _/ w ..d /
•• f ' - Volto,.ru
• - 7J3J 5 - Vold•moro.s I A
• • 1368 6 .. Fronz:.nsbod
---' 7 • Lt:In~nmooDn
07
L
L I) u
., \7 mm
l6 \7 11mm " "
" " " "
the Bulimina-Bolivina-zone (late Badenian) covered the shore of Jurassic limestone. In
the area of the limestone quarry of Devfnska Nova Ves vertical karst fissures are
exposed (Spalte 1-3) with rich large and small mammalian fauna (Zapfe, 1949, 1960,
1979; Fejfar, 1974); the fissures are filled in the upper parts with fine grained mica-
ceous sands without fossils, probably relics of the transgressions. Their lower part
contained mammalian remains in orange and red clays.
Principally, the karstic solution and the filling of fissures precedes in time the
middle Badenian (Lagenid zone) transgression, and could be theoretically lower
Badenian or older. Cicha et al. (1972) and Fejfar (1974), on the base of further collec-
tion in the Neudorf fissures, confirmed correlation with Sansan in the zone MN 6, as
originally suggested by Mein (1979) in defining the MN zones. All important genera in
the assemblage provide (in comparison with similar MN 5 record in Franzensbad,
Langenmoosen, and Maj?endorf) subsequent enlargement of size and gradual changes in
morphology; this concerns especially Eumyarion, Anomalomys, Neocometes, Mega-
cricetodon, and Democricetodon (figures 9-13).
Characteristic taxa of rodents in karst fissures (Spalte 1-3) are (number of

molars given in parenthesis) Eumyarion latior (80), Neocometes brunonis (131),
Anomalomys gaudryi, Keramidomys carpathicus (377), Eomyops catalaunicus (1),
Democricetodon vindobonensis (40), Megacricetodon schaubi (8), Myoglis meini,
Paraglis astaracensis, Pseudodryomys hamadryas, and Eomuscardinus sansaniensis.
The younger second marine transgression of the Bulimina-Bolivina zone is

exposed in its coarse sandy littoral facies in a nearby (Z km to the east of the lime-
stone quarry) sand pit Sandberg, a well known locality (F;.aciostrato~type: Papp et al.,
1976) of late Badenian molluscs and other invertebrates (Svagrovsky, 1974); this is the
site of the marine and land large mammals, including hominids (Thenius, 195Z). This
faunal assemblage is younger than that of the fillings (Spalte 1-3); the differences are
due to another biotope. Both faunas can belong to the zone MN 6.
In 1984, speleologists from Bratislava discovered another rather peculiar site at

Neudorf, also in the karstic fissure system in the Jurassic limestone. The locality
called Bonanza (Holec et al., 1987) is related to the previously known fissure fillings;
however, some layers (coarse phosphatic sandstones) are bedded, and contain on the
surface skeletal units of land and marine mammals (e.g., Pristiphoca, Deinosorex,
Eumyarion) and marine fishes. Thus, the new fissure filling might represent an unique
deposit of a cave under sea level.
Early late Miocene, PaDDonian assemblages of the Vallesian zone MN 9 (or

MN 10). Karst fissures at Suchomasty in Devonian limestones exposed at Cerveny
1om (red quarry), middle Bohemia (figure 1).
The assemblage is rich in small mammals; the large forms are poorly recorded:
small cervid (of Procapreolus size), Tapirus sp., rhinocerotid, fragments of (partly
rounded) mastodon enamel, large viverrid; no remains of Hipparion and bovids.
Small mammalian taxa are as follows (number of molars in parentheses):

Plesiodimylus (30), Scaptonyx (3), Galerix (5), Lanthanotherium (11), Dinosorex (3),
Megaderma (15), Myoglis cf, larteti (15), Anomalomys gaillardi (8), Eomyops cf.
catalaunicus (90), Keramidomys cf. mohleri (5), Democricetodon nov. spec. (167),
Eumyarion cf, leemani (6), Kowalskia cf, fahlbuschi (Bachmayer and Wilson, 1970) (69),
Parapodemus cf. lugdunensis (Z), Protozapus intermedius (16), Glirulus cf, lissiensis
(15), Paraglirulus (Z species: 84), Muscardinus (Z species: 140), Glis sp. (57), Eliomys cf.
assimilis (4), Graphiurops austriacus (1).
In the fauna (figure 14) the glirids are most prevalent with 45% abundance; other
groups and percent abundance: Cricetids (35%), Eomyids (13.5%); Sciurids
(Spermophilinus and two types of petauristids), Dipodids, Murids, and Castorids are
very rare.
227
co
"'"'
2
19
-~- ~
24 25
~
13 14 15
~ IJI 16
w WI 0 1 2 3
\8 1.1 ...
- m ....... \
' I w
m1 /-\ m- ,' I
/ \
2 ' I / \
/ 1 '-' / I ... m! "'
'·' ' I
/ I
/ I / I
/ / I I
,.,. I' '
/ /
' / I /
/ /
I / I / I I
I / / / I
, ___ ........ /
' ....... .... / I /
/ /
... . . _.... ,""' 1.2 I /
(_ ... ~9~Y.l '·'
..•
.
+
·-·~0 ·-·
~·~~ L " L
••
/!!) L
2.o mrn ... ... , ..... ..• 2omm
'·' ·-· " "
Fig. 12. Measurements and represntative record of the genus Anomalomys (Gaillard, 1900). M/1 (1-10), M/2 (11-18), M1 / (19-30). 1-3,
13-15, 19: Anomalomys aliveriensis (Hofmeijer and Bruijn, 1985) from Aliveri; 4-7, 16-17, 20-23: Anomalomys minor (Fejfar,
1972) from Massendorf (4-7, 22, Z3) Rembach (16), Franzensbad (17, 20, Z1); 8-lZ, 18, 24-27: Anomalomys gaudryi (Gaillard,
1900) from Anwil (8, 9, 11, Z7), Devfnskii Nova Ves, Spalte 1 (10, 25), Spalte Z (Z4). Grosslappen-Aumeister (1Z, 18, Z6, 19); 28,
30: Anomalomys gaillardi (Viret and Schaub, 1946) from Montredon.
9 10
2
II I I .I
I w
0 1 2
mm
m.l
w
b
b b-~~
mu 1hudfJ!ff 1.1
mT a-~~
"~ Crhrbhofrn
• 1 - ~ Jifffl:n/ls,
· - ~~~~F.
••• • ~t1Jitlbu'uJJ.1966
L
L 1.1mm
.. u mm "
Fig. 13. Measurements and representative record of M/1 (1-8) and M1/ (9-16) of the genus Neocometes (Schaub and Zapfe, 1953). 1, Z,
9, 10: Neocometes similis (Fahlbusch, 1966) from Dolnice Z (1) and Dolnice 1 (9), Massendorf (Z, 10), and Franzensbad (11,
1Z). 3-8, 13-16: Neoeoiiietes brunonis (Schaub and Zapfe, 1953) from Devfnska Nova Ves, Spalte 1 (3, 5, 14) and Spalte Z (4,
,..,,.., 6-8, 13-16).
<D
~
~2
I
~
\§)
5
• @~
'R
~
6
~'J!.c.:.:;..
'''
17
~8~ 19
Fig. 14. Vallesian assemblage of rodents of the MN 9-10 zone from

Suchomasty, middle Bohemia - I. 1-3: Cricetidae aff. Pseudo-
meriones gen. et sp. indet., right M/1 (inner [ 11 and outer [ 3] view); 4,
5: Eumyarion leemani (Hartenberger, 1965), left M1/, left M/1-Z; 6:
Anomalomys gaillardi (Viret and Schaub, 1946), left M/1-Z; 7-9:
Kowalskia cf. fahlbuschi (Bachmayer and Wilson, 1970), left M/1 (7,
8), left M1/ (9); 1Q-13: Democricetodon nov. spec., left M/1 (1Q-1Z),
left M1/ (13); 14, 15: Paraglirulus nov. spec., left M1/ (14), left MZ/
(15); 16, 17: ~oglis cf. larteti (Baudelot, 197Z), left M/1 (16), left
M 1/ (17); 18: uscardinus d. davidi (Hugueney and Mein, 1965); 19:
Muscardinus cf. vireti (Hugueney and Mein, 1965), left M/1.
The group of glirids is composed of Muscardinus (43.5%), Paraglirulus (Z6%), Glis

(17.5%), Myoglis and Glirulus (4.5%); Eliomys and Graphiurops are rare. The assem-
blage of prevailing glirids (Muscardinus-Paragl irulus-Glis) indicates forested country
with shrubs, baianced with an open savannah-like landscape.
The fauna of Suchomasty is comparable with (1) older assemblages of the late
Astaracian, MN 9 zone of the upper Bavarian molasse, e.g., Hammerschmiede. This
otherwise very similar fauna differs in the absence of murids and early Kowalskia and
by the presence of Megacricetodon; some glirids, e.g., one large specimen of
Paraglirulus is more advanced in comparison to P. assimilis from Hammerschmiede.
(Z) Younger faunas of the early Turolian of middle Europe (e.g., Eichkogel, Kohfidisch,
and Dorn-D~kheim) of the MN 11 zone. They differ from Suchomasty in having more
frequent murids, less diverse glirids (along with the absence of Myoglis), by more
advanced Kowalskia and absence of Democricetodon and Anomalomy gaillardi. The
assemblage of Suchomasty is best affiliated with the poorly recorded levels of
Vosendorf, Geiselberg, and Goh;endorf in Austria, and with the rich fauna from
Rudabanya (Kretzoi et al., 1974) in Hungary. The last fauna .provides a good analogy
in cricetids; the composition of glirids in Rudapanya (most prevalent are Myoglis and
Glis) and rare occurrence of Eomyids are influenced by differences in ecology (more
wet, dense forest).
230
I 3
Suchomostx
Rodentia (n=708)
1 Gliridoe 45%
2 C ricetidoe 35%
3 Eomyidoe 13,5%
4 Sciuridoe 3%
5 Dipod idoe 2%
6 Muridoe 0,1%
7 Casto rido e 0,1%
Fig. 15. Vallesian assemblage and relative

abundance of rodents of the MN 9 or 10
zone from Suchomasty, middle Bohemia-
n. 1, Z: Parapodemus cf. lugdunensis
(Schaub, 1938), left Ml/, left MZ; 3:
Protozapus intermedius (Bachmayer and
Wilson, 1970), left M/1; 4: Eomyops
catalaunicus (Hartenberger, 1967), left
M1-Z/; 5: Keramidomys cf. mohleri
(Engesser, 197Z), left M/Z-3; 6-8:
Para~lirulus lissiensis (Hugueney and Mein,
1965~ left M1/; 9: Graphiurops austriacus
(Bachmeyer and Wilson, 1980), left M1/;
10: Eliomys cf. assimilis (Mayr, 1979), left
M/1.
Probable age of the Suchomasty fauna is MN zone 9 or lower 10.
Early Pliocene, Ruscinian assemblage of the MN 15b zoue. The karst fissures A
and B is Ivanovce, western Slovakia (Fejfar, 1961, 1970; Fejfar and Heinrich, 1985a).
The fauna contain also large mammals, both mastodonts Anancus arvernensis and
Mammut borsoni, Tapirus arvernensis, Parailurus, Parabos boodon, and small cervids.
Better recorded small mammals include Mimomys (Cseria) gracilis, Mimomys
(Mimomys) occitanus, Germanomys weileri, Trilophomys schaubi, Trilophomys
depereti, Kowalskia intermedia, Pliopetaurista, Rhagapodemus, Pliopentalagus,
Hypolagus, Beremendia, Blarinoides, Allosorex (selection of important taxa).
The evolutionary level of both Mimomys species indicate the younger phase of
the MN 15 zone; the enamel crown base of the lophodont-prismatic molars is very low
undulated, and the enamel synclines are without cementum. The assemblages of
different fissures of Ivanovce are temporally closely related and compare best with
231
the faunas from Weze in Poland, Csarn6ta Z in Hungary, and Sete in France (Fejfar
and Heinrich, 1983).
Middle Pliocene, early Romauiau, early Vill8nyian asaemblage of the MN 16a

zoue. Stratified deposits of a volcanic maar (tuffs, tuffites, fine grained sands)
(Fejfar, 1961, 1964; Fejfar and Heinrich, 1985a). Large mammals are similar to those
from Ivanovce but much better represented: Anancus arvernensis, Mammut borsoni,
Tapirus arvernensis, Parailurus hungaricus, cervids (no bovids), Dicerorhinus
jeanvireti. Arvicolids of the genus Mimomys are more advanced; enamel synclines
contain cementum and the crown base of the prismatic molars is more undulated:
Mimomys (Cseria) stehlini, Mimomys (Mimomys) hajnackensis. Microstructure of the
enamel is more differentiated (Fejfar and Heinrich, 198Z). Close affinities of
Mimomys species occur in Arondelli in northem Italy (Michaux, 1971), in Beremend,
southern Hungary, and in Moreda Z and Orrios 5 in Spain.
Latest Pliocene, late Romauiau, late Vill8nyian asaemblages of the last MN 17

zoue. Colluvial debris marls at Ctm~ves, middle Bohemia, karst fissures in P•esivec
and Yc~lare, south Slovakia, in Kolihany, middle Slovakia. These faunas are usually [as
in other sites in Hungary (Villany 3), Poland (Kadzielnia), and Germany (SchemfeldB in
characteristic red soils. Some of the sites are stratified (e.g., VCelare) (Fejfar and
Heinrich, 1983; Fejfar and Horacek, 1983; Horacek, 1985).
The younger mammalian Tertiary assemblages are characterized by abundant

diversified arvicolids - with the majority of taxa with rooted molars (especially
Lemmus-SY(Mptomys and the earliest rootless lagurid Lagurodon). Large species of
Mimomys imomys) - pliocaenicus and osztramosensis - occur together with
smaller forms of Mimomys (Cseria) (e.g., M. reidi, M. pitymyoides, M. tomensis, and
M. stenokorys); also the following taxa are frequent: Villanyia, Ungaromys,
Clethrionomys, Borsodia. The partially stratified sediments of V'Celare (6, 7, 5, 3,
3/B 1) represent the MN 17 assemblages; further sites here (6/8, 6/5, 4/A, 4/D, 1)
belong to the transition to the younger -early Pleistocene- mammal age Biharian,
and are very important for the recognition of the Pliocene-Pleistocene boundary.
The presence of the rootless genus Microtus (Allophaiomijs) (e.g., VC:elare 5)

represents an important marker for the beginning of Pleistocene Fejfar and Heinrich,
1980, 1981, 1983; Horacek, 1985).
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236
THE "PROBOSCIDEAN DATUM EVENT:• HOW MANY
PROBOSCIDEANS AND HOW MANY EVENTS?
Pascal Tassy
Laboratoire de Paleontologie des Vertebres

et de Paleontologie Humaine
Universite P. & M. Curie
4 place Jussieu
75Z5Z Paris Cedex 05, France
INTRODUCTION
The expression "Proboscidean Datum Event" has been coined by Madden and Van
Couvering (1976) to name the major dispersal event which concerned mammalian
faunas and which was known to have occurred in the early Miocene from Africa
toward Eurasia.
It has been known for a long time that the proboscideans ("mastodonts" and
deinotheres) occur abruptly in the early Miocene continental layers of Eurasia. In
Western Europe, the mammalian zone MN4 (now partly MN3b; Mein, 1979) was defined
by Mein (1975) on the basis of the occurrence of proboscideans, namely Deinotherium
and Gomphotherium, together with other new immigrants like the antelope Eotragus
artenensis and the micrommals Megacricetodon and Democricetodon. Subsequently,
the "Proboscidean Datum Event" was situated at 17.5 Ma by Van Couvering and
Berggren (1977). In the same prospect, Aguilar (198Z) defined his zone C 1 by the
immigration of Democricetodon, Megacricetodon, Cricetodon, and of the Proboscidea
at 17.5 Ma. In the interval, Ginsburg (1974, 1979) dissociated in the Loire basin the
immigration of Gomphotherium (at Artenay = MN3b zone) and of Deinotherium
(Baigneaux = MN4a zone). Tassy (1977) retained at Artenay (Loire basin) and Marsolan
(Aquitaine basin), both MN3b localities, two associated elephantoid species, one
gomphothere: Gomphotherium angustidens, and one mammutid: Zygolophodon
turicensis. Hence, the "Proboscidean Datum Event" in Western Europe appeared to be
in fact an "Elephantoid Datum Event."
More recent investigations on Neogene proboscideans tend to somehow compli-

cate the picture (Tassy, 1983, 1985b, 1986). Several elephantoid species which belong
to four monophyletic groups appear in the early Miocene of the Old World. These
groups are: (a) the Mammutidae (true mastodons), (b) the Amebelodontidae (shovel-
tusked mastodonts), (c) the genus Choerolophodon (pig-toothed mastodonts), (d) the
Elephantidae and their paraphyletic stem group, the gomphotheres (including the
Stegodontidae). Reevaluation of the species Gomphotherium angustidens, formerly
described all around the Old World as soon as early Miocene has excluded it from
MN3b zone. Moreover, it has been hypothesized that the species Hemimastodon
crepusculi from the Bugti Hills (only two M3/'s) could represent the ancestral morpho-
type of the Elephantoidea (Tassy, 198Z). It is then crucial to the understanding and
precise definition of the so-called "Proboscidean Datum Event" to identify to which
groups belong the earliest proboscideans from the Miocene of Africa and of Eurasia.

Edited by E.H. Lindsay et al.
Plenum Press, New York, 1990 237
Such an identification is made difficult by the scarcity of the early elephantoid
material. Nevertheless, it is the condition to clear up the pattern of elephantoid
differentiation and the role of the closure of the Tethys Sea.
The Bugti fauna in Baluchistan was the basis of Madden and Van Couvering's
"Proboscidean Datum Event," a fauna supposed to depict the earliest record in Eurasia
of the African invasion, before a wider Eurasian dispersal during the Burdigalian.
Proboscideans listed by Madden and Van Couvering (op. cit.) were: "Gomphotherium
angustidens, G. crepusculi, Zygolophodon tapiroides, Prodeinotherium bavaricum."
Madden and Van Couvering's idea was somehow reminiscent of that illustrated by
de Beer (1964): Southern Asia was the place of a first dispersal from Africa before a
second northern dispersal. More recently, Bernor (1983, p. 36) provided a more elabo-
rated scheme of this model and explained that his Southern and Western European
Province was "periodically isolated from the extensive African emigration because of
a marine barrier posed by the Paratethys and the remnant Tethys Seas." The Bugti
fauna and especially the status of the species Hemimastodon crepusculi bears heavily
on the hypothesis of diachrony of the "Proboscidean Datum Event" between Europe
and Southern Asia, as emphasized by Bernor et al. (1987).
BEFORE THE NEOGENE
"Proboscideans not only appear to have originated in Africa, but remained iso-
lated there until early Miocene times" (Coryndon and Savage, 1973, p. 1 Z3). This long-
accepted statement still is not refuted but needs a short comment. Recent descrip-
tions of Anthracobune and Jozaria, so-called proboscideans from the Eocene of
Pakistan (West, 1980; Wells and Gingerich, 1983), draw the attention to the possibility
that the origin of proboscideans could have been located in Southern Asia. But these
taxa classified either as anthracobunids or even moeritheres do not show clear-cut
proboscidean characters (Tassy and Shoshani, 1988); they can be classified as
Tethytheria incertae sedis. Nevertheless, even if these Southern Asian animals are
the sister group of other proboscideans, they are not Elephantoidea, and have no
influence on the problem of the pattern of elephantoid differentiation.
In Africa true proboscideans (Numidotherium) are known as soon as late early

Eocene (datation given by the charophytes) at the locality of El Kohol in Algeria
(Mahboubi et al., 1986). Other Paleogene African taxa, Moeritherium, Barytherium,
Palaeomastodon, and Phiomia, are known from late Eocene and early Oligocene North
African sites, particularly the Fayum Depression in Egypt. It is known that part of the
Oligocene fauna of the Fayum (Jebel Qatrani) has an Eurasian origin. One marsupial is
an immigrant from Europe (Bown and Simons, 1984), while one tarsiiform primate is an
emigrant from Eurasia (Simons and Bown, 1985). The Asian Paleocene genus
Phenacolophus could be the sister group of embrithopods according to McKenna and
Manning (1977). Moreover, a primitive embrithopod, more primitive than the Fayum
genus Arsinoitherium, has been found in the Eocene of Romania (Radulesco et al.,
1976) and Turkey (Sen and Heintz, 1979). A contact between Africa and Eurasia is
then to be considered to explain the composition of the Jebel Qatrani fauna. Such an
even earlier exchange is also postulated by Mahboubi et al. (1986) and Hartenberger et
al. (1985) to explain the composition of mid-Eocene Algerian and Tunisian faunas.
Yet no proboscidean taxa known in the Fayum Depression and contemporaneous

African sites have been found outside Africa.
THE EARLIEST MIOCENE PROBOSCIDEANS: THE STRA11GRAPBIC RECORD
Europe, Northern Asia
In Western Europe the succession of faunal emigrations is well established with

good correlations between localities from the Loire and Aquitaine basins in France and
from Lisboa area in Portugal (Antunes et al., 1973; Ginsburg, 1974). The arrival of
238
Anchitherium, Stephanocemas, and Brachyodus (MN3a zone) preceded that of
proboscideans. Earliest proboscideans (MN3b) are elephantoids, Gomphotherium and
Zygolophodon. While only the former is present in the fauna RZ of Lisboa, both taxa
are associated at Artenay (Orleanais Sands). They are followed by deinotheres (MN4a
zone: Baigneaux, Chevilly, La Romieu). Proboscideans do not occur in zone MN3a
(Estrepouy, Chitenay in France), which has been correlated with the R1 fauna of
Lisboa (Antunes et al., 1973; p. Z313). This fauna is preceded at Lisboa by the trans-
gressive episode C1 dated (glauconites) Z4 ± 1 Ma (Antunes et al., op. cit.). One
elephantoid molar is alleged to come from an early Orleanian locality (Condom,
Aquitaine basin) and is cited as a possible early occurrence of elephantoid in the
Estrepouy zone (MN3a) (Ginsburg, 1974; Bernor, 1983). This molar (a right MZ/) has
not been found in situ; its precise origin remains unknown. Moreover, its morphology
(a complicated bunodonty with well developed posttrite conules) matches best with
that of Archaeobelodon filholi, a species which makes its first appearance in Western
Europe in the Aquitaine basin, at Bezian (MN4a zone) (Tassy, 1984). It is unlikely that
this molar could represent an early occurrence of this species in Western Europe since
early amebelodontids in East Africa (Rusinga, Set Z of Pickford's (1981) biostratig-
raphy) show a less complicated tooth morphology (Tassy, 1986). Recent careful exca-
vations at Estrepouy by C. Bulot gave no proboscidean remains. The earliest
elephantoid remains in the Aquitaine basin come from Navere (Gomphotherium),
associated with Brachyodus onoideus (Baudelot and Crouzel, 1976), an association
which defines the MN3b zone.
Elsewhere (Eastern Europe, Northern Asia) the early Miocene mammalian faunas
do not show such a detailed sequence, probably due to the scarcity of the findings. In
the East Paratethys, according to Gabunia (1979), among "anchitherian faunas"
elephantoids antedate Anchitherium, a reversed sequence due probably to isolated
discoveries. An early record of "Gomphotherium aff. cooperi" in the Sakaraulian is
correlated with MNZ zone on "stratigraphical position" according to Gabunia (1979, p.
413) and not faunal analysis. Such a datum appears to be insufficient to assess a pre-
Orleanian occurrence of elephantoids in that part of Eurasia. Nevertheless,
Gomphoterium and Zygolophodon, associated with Stephanocemas (Sakaraulian) pre-
ceded deinotheres which appear later (Karaganian), associated with Gomphotherium
angustidens. A more Eastern record is Gomphotherium inopinatum from the Djilancik
Beds in Kazakhstan. Two characters [twisted enamel band on the upper tusks, piri-
form transverse section of the lower tusks with a ventral sulcus, according to
Borissiak and Beliaeva (19Z8)], put this species closer to Gomphotherium angustidens
though not necessarily conspecific.
The Burdigalian fauna of the Hiramaki Formation (Mino, Japan) is the most
Eastern record and shows the association of a gomphothere (the primitive species
Gomphotherium annecteris), with Anchitherium, and Brachyodus (Takai, 1954), an
association known from MN3b zone in Western Europe.
No radiometric data are known for the earliest proboscideans from these areas.
The only record, which came from northern Hungary, cited by Rogl and Steininger
(1983) and by Thomas (1985) can be discarded. Footprints allocated to proboscideans
have been recorded at Ipolytamoc in the "Lower Rhyolite tuff" dated 19.6 ± 1.4 Ma
(Hamor and Ravaz-Barayai, 1979). New examinations of the footprints by Kordos
(1983) prove them to belong to a rhinoceros. The Ipolytamoc fauna is probably older
than MN3b zone.
Southern Asia
There is a general agreement that the Bugti fauna in Baluchistan where

elephantoids and deinotheres are associated is older than the Chinji fauna (Pilgrim,
1908; Raza and Meyer, 1984; Barry et al., 1985; Flynn et al., 1986). Indeed, no Chinji
elephantoid taxa are known from the Bugti Beds which brought Hemimastodon
crepusculi, Gomphotherium cooperi, and Choerolophodon palaeindicus. Taxa from the
Kamlial Formation are less well identified (see revised systematics of Southern Asian
elephantoids in Tassy, 1983). Nevertheless, near Chinji village, Barry et al. (1985)
239
found the earliest gomphoteres and deinotheres (gen. and sp. indet.) from the Kamlial
Formation at the basis of the Chita Parwala-Gabhir section, dated 18.3 Ma (magnetic
polarity) by Johnson et al. (1985). They conclude that the Bugti fauna must be older.
The question of the correlation of the Bugti fauna with other early Miocene localities
is much debated and Pickford (1987) equated Bugti with Maboko in East Africa (Set 3,
i.e. circa 15-16 Ma).
Africa
The earliest dated African Neogene proboscidean is a mammutid, Eozygodon

morotoensis found at Meswa Bridge, Kenya (Pickford and Tassy, 1980; Tassy and
Pickford, 1983). The locality is dated (biotites) between Z1 and ZZ Ma (R. Drake,
unpublished). This locality antedates the classical early Miocene East African
mammalian localities of Set 1: Koru, Songhor, Legetet (Pickford, 1981). This East
African species is known only at Meswa, Moruorot, and Moroto. The two latter local-
ities are also of early Miocene age (Set 1). Moroto in Uganda (type locality) was
situated early or middle Miocene (Pickford and Tassy, 1980; Tassy, 1986) but new
finding and faunal studies at Moroto show it to belong to Set 1 (Pickford et al., 1986).
Deinotheres are not known at Meswa but this can be due to taphonomic or other
factors; only one partial skeleton and a deciduous tooth of Eozygodon morotoensis
were recovered at Meswa. Other elephantoids of Set 1 and Set Z (cf. Archaeobelodon,
Archaeobelodon aff. filholi, Gomphotherium sp., according to Tassy, 1986) are associ-
ated with deinotheres.
The Arabian PeoiDsula
Northern localities in Yeroham and Rotem basins (Negev) and southern localities
of Ad Dabtiyah and As Sarar (Saudi Arabia) yielded early Miocene faunas. Though the
Negev faunas have been compared with "late MN3a" (Tchernov et al., 1987), a correla-
tion with MN4-MN5 zones is likely (Tassy and Thomas, 1988). Prodeinotherium is
associated there with Eotragus cf. sansaniensis, cf. Canthumeryx, and a hypsodont
gazella. Elephantoid remains are scarce but not particularly primitive. Saudi Arabian
localities could be of the same age, i.e. "MN4b" (Thomas, 1985), or "mid-Qrleanian"
(Whybrow, 1987). Gomphotherium sp. is cited at As Sarrar (Thomas et al., 198Z) and
Gomphotherium cooperi is cited at Ad Dabtiyah (Gentry, 1987). This latter assign-
ment does not necessarily mean that Ad Dabtiyah is as old as Bugti from where G.
cooperi was first described. The primitive bunodont pattern of some molars from Ad
Dabtiyah fit well with that of molars of the Gomphotherium "annectens group" [i.e. G.
cooperi but also G. annectens from Japan, G. sylvaticum from Western Europe and
Gomphotherium sp. from Mwiti and Buluk in East Africa (Set zll. There is a high
measure of variation in the Ad Dabtiyah collection and perhaps two elephantoids are
present. A flat lower tusk (Gentry, 1987, fig. 17) matches best with that of
Archaeobelodon a££. filholi, an amebelodontid described by Tassy (1986) at Mwiti and
Buluk (though it must be kept .in mind that the flat transverse section is a primitive
character). Lower tusks of the G. "annectens group" have piriform to oval transverse
sections but lower tusks are not known at Bugti.
EARLY MIOCENE PROBOSCIDEANS: SYSTEMATICS,

RELATION~,ANDPALEOmDGEOGRAPHY
Deiootheriidae
On the basis of the stratigraphical record, the first deinotheres (Prodeinotherium

Ehik, ~Harris, 1978) followed the elephantoids in Western Europe. This is proba-
bly indicative of diachrony between the arrival of elephantoids and deinotheres in this
area. As far we know, deinotheres do not occur in Northern Asia. Because of the
numerous known Miocene mammalian remains from China, where deinotheres are
always lacking, it is very unlikely to anticipate future discoveries of deinotheres in
this region. Perhaps this situation is due to ecological constraints, deinotheres being
inhabitants of rather forested environments. Yet, a detailed analysis of the different
240
Ha 2
j
·~ ..,c
0
·~
Chios, Soft;a
15 AS 6
~ "
0
·.:;~ "ll
~ ~
16 ·~
"
~
~ ~
.il ~ ~
"c
17 ~
~
~ "'
0
0
~
0
] Bezian
'6
·-~,;; {l"
18 ~
Artenay
Rusluga (HJwcgl) .l!
2 : t;
Hfwangano ll
19
J.egetct " '
' ll ~
20 ~ ~
"'
+-"--
1 Bugt 1
21
Meswa
22.
A B
Fig. 1. A: Stratigraphic position and relationships of the earliest members of the
four elephantoid monophyletic groups and of their putative "ancestor" H.
crepusculi. The Bugti Beds are hypothetically situated at 20.5 Ma.
B: Stratigraphic position and relationships of the earliest members of the
four elephantoid monophyletic groups in Western Europe. [1: mammal ages
(Fahlbusch, 1976), Ag = Agenian, 0 = Orleanian, As = Astaracian;
2: mammal zones (a: Mein, 1975; b: Mein, 1979)1.
species allocated to Prodeinotherium, P. pentapotamiae in Southern Asia, _E. hobleyi in

Africa, and _E. bavaricum in Europe, is needed to give light on the pattern of the early
differentiation of deinotheres.
E1ephantoidea
The mammutid species Eozygodon morotoensis is the earliest dated Miocene

elephantoid (circa 22 Ma), while the most primitive known Neogene elephantoid,
Hemimastodon crepusculi, has been recovered from the Bugti Beds. As far as terres-
trial animals are concerned, the hypothesis of a contact between the Afro-Arabian
plate and Eurasia before 22 Ma (figures 1 and 2) is supported by sister group relation-
ships of Hemimastodon crepusculi with other Elephantoidea, its occurrence in the
Bugti Beds, and by the sister group relationships between Mammutidae on the one hand
and all other elephantoid groups on the other (Tassy, 1982, 1985a,b).
The status of Bemimastodon crepusculi (Pilgrim). I considered Hemimastodon

crepusculi a proboscidean, and the sister group or ancestral morphotype of the other
Elephantoidea (Tassy, 1982). According to this view, only two left M3/'s were allo-
cated to that species (one of them is the holotype). If H. crepusculi is the sister group
of the Elephantoidea (including Mammutidae), the date of the differentiation of the
two sister groups must be older than 22 Ma, date of the first appearance at Meswa
Bridge of the mammutid E. morotoensis. The biostratigraphic consequence is that the
"Proboscidean Datum Event" could be at least as old as 22 Ma. Then, the differentia-
tion of Elephantoidea could have occurred in both Africa and Southern Asia. Conse-
quently, part of the Bugti fauna could be older than other known Eurasiatic faunas, or
contain a relict primitive elephantoid species, evidence of an earlier dispersal, yet
with very few fossil records (though an early marine disconnection was postulated by
Drooger (1979) about the end of. the Chattian on the basis of larger foraminifera).
However, the systematic assignment of the Bugti species has long been discussed
and is still debated. It is either a primitive "mastodont" (Pilgrim, 1912; Matsumoto,
1927; Hopwood in Osborn, 1936, p. 236; Chakravati, 1957; Tassy, 1982), or allocated to
Gomphotherium (Madden and Van Couvering, 1976), or even synonymized with G.
241
,~·
•
~
\
\ \
\
\
\
\
\
\
\
\
\
\
\
\
\
\
\
\
\
eo
1~
2•
3+
4•
so
6~
Fig. Z. Differentiation of the Elephantoidea: distribution of the earliest members

of the main Neogene monophyletic groups and of their Paleogene sister
groups. 1: Phiomia (Oligocene, Jebel Qatrani, Fayum); Z: Hemimastodon
crepusculi (early Miocene, Bugti, Baluchistan); 3: Eozygodon morotoensis
(early Miocene, Meswa, Kenya); 4: cf• .Archaeobelodon [early Miocene,
Legetet, Rusinga (Kenya)]; 5: Gomphotherium "annectens group" (early
Miocene: Bugti, Baluchistan; Mfwangano and Mwiti (Kenya); Hiramaki Fm.,
Japan; Artenay (MN3b zone), France; Qta. das Pedreiras (MN4a zone),
Portugal); 6: Choerolophodon (early Miocene: Bugti, Baluchistan; Buluk,
Kenya). Map: "Dymaxion" ground-plan, by R. Buckminster Fuller.
242
Ma
4
. ..~I
-~I
.8 a:
Series
~ Blancan
::•
... .....
5 • "'"I
6
:• i•a I
.... :1:1
7 li'
T
,&"' Pil<ermi \'? Hemphillien
8
'<;-
\ i.
.i
9 Sob lay
10 v
..."'
Dinotherien-
sande .~ ::larendonien
11 :~lchinji .~
?
\ iii I
12
8 ~I I ·~ I "
I
Montrejeau ·~ e I "
13 ;
..;1,.1 ~· i...
gI Malartic ~ I·~ I "I
14 AS 7 i 1villefranche"l: I~ 1rung s:: INianr;xia
~
"
i
~
1
Cherichira .a .g·.a I Gur .:t hunnan w
Barstovian
15 6 Ulan ,..1 Sansan.- c.l ., I "i:
..."' Tologoj ..; I •·i! ~I
I .Pontlevoy I l; I
_,..-
16
~=
.. "'6
II
~
17 NORTH AMERICA
~ Moroto Ojilancil< ;~
Beds 1..; Artenay ,Pontigne
18 ~ Moruorot l;.l
l .._,1 Zygo!.ophDdDn
19 ~ ~~nsis group"
20
EAST AFRICA Meswa ~
21
~ NORTH AFRICA
EURASIA
22
Fig. 3. Phylogenetic relationships and chronological range of the Mammutidae.

Thick line: known chronological range; dash line: possible extension.
angustidens (Sarwar, 1977); the holotype is considered an abnormal tooth (Schlesinger,

1917; Cooper, 19ZZ) or supernumerary (Tobien, 1973). On the other hand, it is assigned
to "a suoid or a bunodont anthracothere" (Matthew in Osborn, op. cit.) or precisely to a
suoid, on the basis of the triangular shape of the crown in occlusal view (Osborn, op.
cit.). The suoid hypothesis has been renewed in more detailed terms by Pickford
(1987). According to this hypothesis, the question of a Bugti "Proboscidean Datum
Event" earlier than the European "Proboscidean Datum Event" is no longer to be
discussed. I summarize here the arguments developed elsewhere (Tassy, 1988) to
maintain the proboscidean interpretation.
Pickford (1987) described in the Bugti Beds a giant kubanochoerine on the basis
of lower dentitions and allocated to this suoid the holotype of H. crepusculi and the
other M3/ described by Tassy (198Z). Pickford did not discuss the characters of the
M3/ I used to place H. crepusculi in an intermediate position between the Fayum genus
243
Ma
~ ~
z
w
:E
.
.! i-
4
R .~
21
IS
}1.
5
r ll ] .g
6 ] ~ ~
eo"
"'~"" l'laraghen i
Kertch '!' ~
Sahabi ~ Rhino Hi 11
'i
7 Q.
8 T Ohok Pa than 1 j"

~ Mixson
~
?
"' "
,Q
.3' Snake Fm
~I Xmas Quarry ~II
10 v t I
·=~ "''?
I
11 ... :::, Chinji
~ NORTH AMERICA
"r ~
~
12
13
8
La Grive 'i ..;
;
.·;::
AS ;!: il Tung Gurl

·~t
14 ~ For} Ternan
_...
;::~ Ningxia 1 ~
15 6
.i ...
«:!. ~
:! ;:~rri sdri ft
~ • ~aY~
/';. 0 ~0k 1 · ,-·'
·~~~Aropl
:·· . J.:f ..... :Sind
Sansan • uru a 1 i
16 :::: Bi rosse ~~Be l:ometches kay a
17 i ~
Hwitil sezian
J
......... Lopero.!,t._)
·-..,,-·'
0 ]
Rusinga a BulukJ • __ ... --- .--~-·-·!
18
12 · ...~: WESTERN EUROPE
·--··
~
19 ... : Jl
Legetet .... ~
20
... -·· .... -- .--' .!l
ll.
21
22
Fig. 4. Phylogenetic relationships and chronological range of the

Amebelodontidae. Thick line: known chronological range; dash line:
possible extension.
Phiomia and Elephantoidea. On the contrary, he listed five characters said to be

kubanochoerine characters. Four of them are known to be regular characters of both
Phiomia and primitive gomphotheres or other Neogene groups (thick cingulum,
trilophodonty, narrow distal part, weak "conelets," transverse lophs in shape of an
open V). The last character (shape of the roots) does not match with my observa-
tions. Nevertheless, the striking features of the two M3/'s, i.e., trilophodonty ("talon
bicuspide") and strong lingual cingulum, are used by Pickford (1987, p. Z98) in the
diagnosis of the genus Hemimastodon among Kubanochoerinae. In the subfamily, the
M3/ is bilophodont with a strong disto-lingual cusp but there is no labial cusp (i.e., no
third loph) and the labial cingulum is strong, although not the lingual one, even in the
biggest known kubanochoerine from Africa (Wilkinson, 1976, p. ZZ7, Z3Z). I add that in
kubanochoerines the labial cusps are rounded, not transversally elongated as in H.
crepusculi and other elephantoids. .The M/3's allocated to H. crepusculi by Pickford
have a typical suoid morphology. The hypoconulid, though massive with a lingual
sulcus does not form a tritolophid (i.e., no symmetrical labial and lingual half-lophids
separated by a deep median sulcus); its morphology is close to that of other known
kubanochoerines. If those M3/'s and M/3's were associated, M3/ would be more
244
derived (in an elephantoid manner) than M/3, an unlikely situation. In proboscideans
and suoids, M/3 is always more complicated than M3/. Also, the two M3/'s are longer
or as long (99 mm and 92+ mm; Tassy, 198Z) as the M/3's (63-98 mm; Pickford, 1987, p.
30Z), which is unusual.
Indeed, there is one giant suoid in the Bugti Beds. The elephantoid features seen
on the two M3/'s could well be due to convergence and linked to a bunolophodont
pattern. This is a possibility I do not underestimate. Nevertheless, on a strict char-
acter analysis, I maintain that the proboscidean hypothesis is not refuted. No char-
acter known only in kubanochoerines and not in Elephantiformes has been described,
based on the M3/'s of H. crepusculi. As I wrote in 198Z, no M3/ of this kind is known
among Artiodactyla. Microstructural analysis of the enamel could give further
information. But the association in a maxilla of such a trilophodont M3/ with
kubanochoerine premolars and molars is needed to warrant Pickford's hypothesis.
The early differentiation of Elepbantoidea. Among Mammutidae (figure 3), the

primitive species Eozygodon morotoensis is endemic in East Africa. A more evolved
species, Zygolophodon turicensis, appears in Western Europe at Artenay (MN3b zone)
while Zygolophodon sp. is cited in the basal Manchar Formation by Raza et al. (1984).
The species Zygolophodon turicensis is frequently cited in the faunal lists of the
European Miocene but its anatomy and evolution are not so well known. Clearly
enough, its occurrence in the European Orleanian postdates the early differentiation
of the Mammutidae.
Among Amebelodontidae (figure 4), the earliest European species, Archaeo-

belodon filholi, appears in the Aquitaine basin at Bezian, near La Romieu (MN4a zone),
associated with Gomphotherium sp. and Prodeinotherium sp. Other Orleanian locali-
ties (MN4, MN5 zones) are Pontlevoy-Thenay and Tavers in the Loire basin, Birosse in
the Aquitaine basin, and Quinta da Noiva, Q. da Narigao, Q. da Farinheira in
Portugal. A. filholi is abundant at Sansan (MN6 zone). The species was formerly
confused with G. angustidens and was recently revised by Tassy (1984, 1985b). A
closely related form, A. aff. filholi has been described (Tassy, 1986) in East Africa at
Mwiti and Buluk (Set Z). On the basis of cranial characters, the East African form is
more primitive than A. filholi at Sansan but such characters are not seen on Orleanian
remains. The lineageis present in East Africa at Rusinga (Kulu and Hiwegi: Set Z)
and perhaps at Legetet (Set 1), i.e., already present between ZO and 18 Ma. But more
material is needed to identify the precise relationships of the early Miocene East
African form and the European Orleanian A. filholi. In Southern Asia no
amebelodontid is recognized in the Bugti Beds:- The group appears in the medial
Manchar Formation of Sind (Raza et al., 1984) where it is allocated to "Protanancus
chin·iensis ou espece proche" by Tassy (1983, p. 1Z8). Perhaps also the specimens from
Sind one partial tusk, two molars) could be allocated to Archaeobelodon.
Among gomphotheres (figure 5), the species Gomphotherium angustidens as

revised by Tassy (1985b) is restricted in Western Europe to late MN5 zone (Le Moune,
Aquitaine basin) till MN9 zone (Eppelsheim). It is widespread toward Turkey where it
is known at Pafalar (Gaziry, 1976). Outside this area its occurrence is dubious.
The early European gomphothere Gomphotherium sylvaticum (Tassy, 1985b)

belongs to the G. "annectens group," a group of species widespread in Eurasia, though
not abundant. This species represents in Europe the arrival of the Elephantoidea
(Artenay, MN3b zone). It was formerly confused with G. angustidens but shares with
other taxa of the G. "annectens group" primitive characters distinct from G.
angustidens. (Tassy, 1985, p. 685). Its occurrence in zone MN4a is only supported by
one specimen from Quinta das Pedreiras (R3 fauna from Portugal). One isolated M3/
is supposed to come from the Falun of Pontlevoy, but the exact origin is unknown; this
is insufficient evidence to cite the presence of G. sylvaticum in MN5 zone. Members
of the "annectens group" are rare in East Africa; Gomphotherium sp. (Tassy, 1986) has
been discovered at Mfwangano (two specimens) and Buluk (one specimen), i.e. Set Z
(circa 18.5 Ma and 16.5 Ma). The relationships between primitive gomphotheres are
still unclear since this is a "grade group." Nevertheless, in Europe this grade clearly
245
(/)
QJ
""QJ
.c
.u
0
9 .c
~
10 v 00
11
12.
8
13
14 AS 7
15 6
16
17
18
19
20
21
22
Fig. 5. Chronological range and relationships of trilophodont

gomphotheres depicting the possible origins of North
American gomphotheres and of tetralophodonts (a group
including elephants and stegodons). 1: Mino, Hiramaki Fm.
(Japan); Z: Artenay, O:~;leanais Sands (France); 3: Mfwangano,
Mwiti (Kenya); 4: Bugti Hills, Baluchistan (Pakistan);
5: Djilancik Beds, Kazakhstan (USSR); 6: Birosse, Gers
(France); 7: En Pejouan, Gers (France); 8: Gebel Zelten
(Libya); 9: Beaugency-Tav ers, Orleanais Sands (France);
10: Eibiswald (Austria), Sandelzhausen, O.S.M. (West
Germany); 11: Montrejeau (France); 1Z: Eppelsheim,
Dinotheriensan de (West Germany); 13: Steinheim (West
Germany); 14: Potwar Plateau, Chinji Fm. (Pakistan). Thick
line: known chronological range; dash line: possible extension.
246
M !e
S!! ~z
:c ;:;::;
u
"'=.....
::::0
:IE
2 ...
:c
C[
4
R
5
7 U. Maragheh I
Samos
8 T Pikermi
.... Dhok Pathan ••
Dj ebel Hamrin .~ Hart Talyangar ~
;"'""
0.:
9
Kayadibi
...=
L.Mara~heh t Nagri
10
v Yassioren t..i
Ravin de la Pluie Nakali
t..i
11 Kiiciikcekmece 2
Esme Ak~ akoy 1 Ngorora 9 Chinji
12
8
C.palaaindiauc/
Akin ~ Beglia
I
l ~ I
I
I
13 kir.umuensis group
I~ Sari~ay ~~~.
?~~-"I
I
...--~"''-----,
r~ Sof~3
14 AS 7
I .,
Chi I'"
os 1 ij EUROPE
I .,.
I
a.l .. lt..i
15 6
Kamlial ~ I ~~ ..aoo
b k
.,<.Jj u
16 5 I ·~
17 i I ~
·s 1Buluk
0 .s
18 a
19
SOUTHERN ASIA
Bugti
20
AG
21
22
Fig. 6. Phylogenetic relationships and chronological range of

Choerolophodon. Thick line: known chronological range; dash
line: possible extension.
247
antedates the occurrence of G. angustidens. I tend to abandon the idea of a derivation
in Western Europe of the latter species from G. sylvaticum via an intermediate form
usually called "G• angustidens subta iroides" (MN4-MN5 zones). A later emigration of
North African origin (Gebel Zelten toward Western and Eastern Europe (G.
inopinatum) can be considered. On the other hand, the derivation from G. sylvaticum
of the rare species G. steinheimensis known at Steinheim (MN7 zone) is better
supported (Tassy, 1985b).
Among choerolophodonts (figure 6), the Bugti Beds yielded the primitive species
Choerolophodon alaeindicus (Lydekker). The species (two molars only) was newly
defined by Tassy 1983, p. 259). Its occurrence antedates the closely related, if not
conspecific, East African species Choerolophodon kisumuensis described at Maboko
(Set 3, circa 15-16 Ma). The diachrony shown by the distribution of the two species
has been used by Tassy (1983, p. 260; 1985b, p. 642) as possible evidence of a late
dispersal ("Neogene Dispersal Phase 2" of Thomas, 1985) from Southern Asia toward
Africa circa 16 Ma. This hypothesis must be abandoned. At Buluk, in a locality older
than 17.2 Ma, according to Leakey and Walker (1985), a "new genus and species of
gomphothere" cited by these authors can be allocated to a primitive Choerolophodon
(Harris and Tassy, in prep.). The stratigraphical gap between Bugti and East Africa is
then shortened. The exchange, still perhaps from Southern Asia toward Africa, could
have occurred during what is usually called the "Proboscidean Datum Event" (circa
18 Ma). A probable Choerolophodon is also "Mastodon spenceri" from Moghara, Egypt
(Fourtau, 1920), but the original material has never been revised.
In Europe the earliest choerolophodont, Choerolophodon chioticus described by

Tobien (1980) at Chios, is of Astaracian age, possibly MN6 zone. Other pre-Hipparion
choerolophodont remains are described at Sof~a by Gaziry (1976). The mid-Miocene
occurrence of the species is evidence of a late main dispersal phase involving
elephantoids known elsewhere from early Miocene localities. Distribution of early
choerolophodonts shows clearly a provinciality between Southern Asia and Africa on
the one hand and the Eastern Mediterranean region on the other (in the latter area the
genus, abundant during the late Miocene, is not known west of Yugoslavia).
This late emigration was already postulated by Adams et al. (1983) to explain the
occurrence in that area of other mammals, Megapedetes, Orycteropus, and
Kubanochoerus. In the Eastern Mediterranean area, another elephantoid, the
amebelodontid Platybelodon danovi, appears probably at the same time (MN6 zone)
though not associated with Choerolophodon chioticus. It is known at Belometcheskaya
(perhaps MN5 zone according to Mein, 1981) and also at Arapli, western Turkey, a
middle Miocene locality according to Gaziry (1976). The earliest record of a
platybelodont is Platybelodon sp., represented by a characteristic lower tusk from the
early Miocene (Set 2) of Loperot, East Africa (Maglio, 1969; Tassy, 1986). It
emphasizes the reality of this late emigration of taxa already present in the early
Miocene of Africa.
CONCLUSION
In Europe, the so-called "Proboscidean Datum Event" is restricted to the

immigration at zone MN3b of Gomphotherium sylvaticum, a gomphothere which
belongs to the G. "annectens group," and of the mammutid Zygolophodon turicensis.
Other taxa known in the early Miocene of Africa and Southern Asia come later:
deinotheres and amebelodontids at zone MN4a (perhaps associated with another
gomphothere ancestral to G. angustidens), and choerolophodonts during the
Astaracian, possible zone MN6.
On the basis of phylogenetic relationships between the main Neogene groups and
of the stratigraphical record, hypotheses of diachrony and provinciality can be
drawn. They are working hypotheses since the fossil record of early Miocene
elephantoids still has to be enriched to grasp the thin resolution we need to solve the
problem.
248
An early contact (22 Ma or more) between Africa and Southern Asia let primi-
tive elephantoids such as Hemimastodon crepusculi differentiate into Mammutidae on
the one hand (perhaps in Africa) and Amebelodontidae, gomphotheres and
Choerolophodon on the other. A subsequent emigration toward Western Europe and
Northern Eurasia corresponds to what was usually called the "Proboscidean Datum
Event" and situated circa 18 Ma [circa 19 Ma according to Rogl and Steininger (1983);
between 18-19 Ma (="Neogene Dispersal Phase I") according to Thomas (1985); "mid-
burdigalian" according to Adams et al. (1983)]. In Europe, this "event" is broken into
three phases, two of them taking place during the Orleanian. The first is the arrival
of gomphotheres and mammutids. The second is the arrival of deinotheres and
amebelodontids. The third, limited to the Eastern Mediterranean region, is later
(Astaracian) and involves the arrival of choerolophodonts together with a
platybelodont.
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252
THE PROBOSCIDEANS DATA, AGE AND PALEOGEOGRAPHY:
EVIDENCE FROM THE MIOCENE OF LISBON
M. T. Antunes
Centro de Estratigrafia e Paleobiologia da UNL

Quinta da Torre
Z8Z5 Monte da Caparica, Portugal
INTRODUCTION
Proboscideans originated in Africa. Mastodonts are descendants of Eocene

moeritheres or related forms. Deinotheres may be descendants of barytheres, known
only from late Eocene and early Oligocene.
In early Miocene, mastodonts and deinotheres migrated into Eurasia. Not much
later, mastodonts were also established in North America.
Expansion in Eurasia was fast. Expansion events are therefore meaningful under
both age and paleogeographical viewpoints.
There is not one "Proboscidean datum," but indeed two. In western Europe,
deinotheres arrived somewhat later. The first proboscidean immigration is thus that
of tnastodonts. How did this situation develop?
Evidence from the Lisbon Miocene sequence is useful because there is a wealth
of correlative data in that area. Paleogeography, climate, and how they affected this
Eurasian land's end (among other areas) during Miocene times are also of interest.
lDSTORICAL APPROACH
Mastodonts have been recognized near Lisbon since Roman (1907, p. 53-54): a
tooth was recorded for the V-.!_ division (local stratigraphical unit), then regarded as
lower Helvetian and now as uppermost Burdigalian. Teeth from later units are not
dealt with here. Stratigraphical setting is that of Cotter (1904), still in use with some
modifications.
The presence of mastodonts in the same levels was discussed by Torres (1935).
Further developments came from G. Zbyszewski, who reported on vertebrate

fossils collected in Lisbon's sandpits (Zbyszewski, 1937). He also recorded the
presence of Deinotherium for the first time in Portugal (Zbyszewski, 1941) based on a
specimen whose origin is not exactly known; however, its patina seems to indicate the
pyrolusite-rich sands from V-.!_Z subdivision near Lumiar, where most of the subsequent
specimens were found. This also applies to one tooth from Quinta das Pedreiras
(Lumiar), wrongly included by the same author (Zbyszewski, 1949, p. 74) among the
specimens he looked upon as from the Burdigalian IV-E. division- but this is certainly

not correct since patina and other data also indicate V-~Z its true origin (30 years of
collecting did not add any further evidence).
A general memoir on Portuguese mastodonts was published some years later

(Bergounioux et al., 1953).
The successive appearances of the first mastodonts (gomphotheres) and

deinotheres were clearly recognized in the Lisbon basin (Ginsburg and Antunes, 1967,
p. 136-137) as well as in the Loire basin, France. They were not synchronous events,
as it would seem previously.
Several contributions to the paleontological, time, and stratigraphical framing

have since been published. For a general view see Antunes (1979, 1984). Paleoclima-
tological data were provided by Antunes and Pais (1983) and by Pais (1986).
PROBOSCIDEAN BIMIGRA110N - L1MlTED IN TIME?
Gomphotheres appear in the Lisbon area for the first time in the IV-b fluviatile
sands (" Areias da Quinta do Bacalhau") (figure 1). However, is it the real first occur-
rence, or could ~omphotheres arrive earlier? The same question applies also to
deinotheres, whose first known occurrence is in V-~Z "Areias com Placuna miocenica."
In both cases there is no evidence of an earlier arrival. No proboscidean remains

older than IV-]!_ have ever been found, in spite of their generally large size and hence
being easy to spot. Indeed, workers have often kept much smaller fossils as fish or
Cainotherium teeth. Deposits older than IV-b beds have also been exploited, even for
commercial purposes, yielding mammalian fossils but not including the faintest trace
of proboscideans. Examples are the II division localities that are rich in lower
Burdigalian small mammals, MN3a zone (Antunes and Mein, 1986). Another example is
the Horta das Tripas site in Lisbon, upper part of I unit, which is lowermost
Burdigalian; an antbracothere, two rhinocerotids, and a few other mammals were
found, yet no proboscidean. It is most unlikely that proboscideans would remain
unnoticed if there were any present. A counter-proof concerns other littoral marine
deposits near Lisbon - gomphothere teeth have been found in nearly all later middle
Miocene units.
Particularly interesting is the arrival of gomphotheres into North America, the

most distant area from its origin. In Pliocene times, mastodonts passed into South
America from North America. Presence of proboscideans has been recorded in North
America since the Barstovian land mammal age, about 16 to 1Z Ma, which corresponds
quite closely to the Langhian and a large part of the Serravallian in Europe (Berggren
et al., 1985). Some slightly different figures are given for Barstovian, 16.5 to about
11.8 Ma (Hulbert, 1988, fig. Z9, p. 31Z), but that does not change anything. This means
that gomphotheres arrived into North America only somewhat later than into
westernmost Europe, or roughly 17 :i: 0.5 vs 16 :i: 0.5 Ma. The entire migration process
took not more than about Z Ma at most, and maybe 1 Ma or less. Diacbrony of the
first arrivals everywhere in their huge distribution area does not seem to be very
meaningful. This contrasts with a reverse migration later in the Miocene, that of
Hipparion into the Old World.
All of this shows that migrations like these happened during a short time span,
provided there were no really impeaching climate or physiographic barriers. We may
assume, therefore, that the Gomphotherium datum and Deinotherium datum are very
good time references in Europe. Yet they are not chronological, ecological, or paleo-
geographical equivalents.
254
.
:;;
~ 15
riiii'7iit Qu inta da Farinheira htuna

-- -~ ~ CHis~anotherium)
z
Ill
~1r"'\ __ ,._IMN4AI Quinta do Pombeiro faunaS
Dein,
0
-- r?).,...IMN3BI Quinta do Nar igio f•\ma~
!NN.ol! ~ Gomph
0
Universidade CatO iica and

- -- - - - - - - - - - ... ~Av . do Uruguay fauna
N•aNS) -- - ------ ~~Horta du Tripas fauna
0 •
- a. 25
.J ..
o=>
Fig. 1. Lithological succession (part) in Tagus basin near Lisbon, showing rela-
tionships between marine and continental sedimentary bodies. Most
facies are marine; littoral, shallow-water biocalcarenites and silts pre-
dominate over conglomerates and fringe barrier, coralic limestones (only
in Aquitanian). Transitional facies include littoral beach and dune sands,
pelites (often in association with oyster banks), pelites with oligohaline
mollusks (as Cerithium), etc. and plants, swamp pelites with lignite, and
lagoon pelites. Nonmarine facies include fluviatile conglomerates and
often feldspar-rich sands. I to VIa = local stratigraphic divisions and
associated fauna. CO to C5 = sedimentary cycles, each corresponding to
a transgression event and next regression (TO to T5 transgressive events,
see text). RO to R4 = maximum regression episodes marked by intensive
accumulation of terrigenous sediments, mostly corresponding to large
river facies. N4 to N13 = stratigraphic position of samples with plank-
tonic Foraminifera (with corresponding Blow zonation) and a K-Ar
glauconitic age (another K-Ar age for the same glauconite is of only
about Zl Ma; however, it seems too young in this context). NN4 and NN6
= calcareous Nannoplankton zones of Martini. - - - > first arrivals of
gomphotheres and deinotheres. MN3a = lower Burdigalian Horta das
Tripas fauna and slightly younger Univ. Catolica and Av. do Uruguay
fauna, both assigned to that mammal units by P. Mein. MN3b =
Burdigalian Quinta do Narigao fauna, yielding the last Brachyodus
(anthracothere) and the first gomphotheres. MN4a = latest Burdigalian
Quinta do Pombeiro fauna with the first deinotheres. MN4b = Langhian
Quinta da Farinheira fauna (Hispanotherium fauna).
AGE: CHRONOLOGICAL FRAMING
Considering the two proboscidean data, marked by the first appearances of

gomphotheres and deinotheres in Lisbon's IV-b and V-a units, discussion will be based
upon biochronological and stratigraphical evid;nce froiii" this region.
- -
IV-b and V-a Mammalian Faunas
Miocene mammalian faunas have been discussed elsewhere (Antunes, 1984).
Fauna from IV~ comprises the last anthracotheres and the first gomphotheres.
No taxa suggest any immigration path into Iberia other than through the region where
255
France is today. Immigrants just joined forms already living there. Among rodents
there is a majority of glirids, but no cricetid. It corresponds to Mein's MN3b zone and
is more or less contemporary to Ateca m in Aragon (Spain) and Artenay in France.
Next, V-.!. fauna mainly differs by the presence of newcomers as Deinotherium

and Gaindatherium (Iberotherium), and by the absence of anthracotheres, locally
extinct by then. Glirids still predominate among small mammals. This fauna may be
assigned to MN4a zone, and is close in age to Valtorres and others in Spain or to La
Romieu in France (Antunes, 1984, p. 310).
Preceding FilUIIU {Upper I and D Units)
The oldest association, Horta das Tripas (upper I), comprises a few early
Burdigalian larger mammals. It is not comparable to the somewhat younger fauna
from Universidade Cat61ica and Avenida do Uruguay (Antunes and Mein, 1986), which
are also MN3a but only with small mammals. It is an approximate time equivalent of
Ateca I and Estrepouy in France (Antunes and Mein, idem). It may also be correlated
to the "Untere Landschneckenmergel" in Frankfurt Nord basin (Tobien, 1987, p. 2.13), a
time equivalent for N 5 (planktonic foraminifera) and NNZ (calcareous nannoplankton)
zones (cf. Steininger et al., quoted by Tobien, id., Tab. 1).
Following •Hispanotherium-F8111Ul• (V-]!_ Unit)
After the brief marine event corresponding to V-.!_3 subunit ("Calcarios da

Musgueira") there has been a regression marked by the large-scale influx of fluviatile
sands (" Areias do Vale de Chelas," V-~ early middle Miocene) yielding a new and quite
different fauna. Among newcomers, the most remarkable element is the Asiatic
rhinoceros Hispanotherium; cricetids predominate over scarce glirids. This regression
can be assigned to MN4b zone.
Planktonic Foraminifera, Ostracoda, and Calcareous Nannoplankton
Interfingering of marine and nonmarine deposits allow direct correlations. In

order to frame the gomphothere and deinothere data, attention must be paid to inter-
calated marine units.
Prior to IV-~ and beginning with the marine littoral sands of unit n, units m
("Banco Real, n mostly biocalcarenites) and IV-.!. schlier facies, pyritous clays (" Argilas
do Forno do Tejolo") are developed, corresponding as a whole to the main Burdigalian
transgression (TZ). This succession ends in confined marine and brackish facies, over-
lain by IV-~ fluviatile sands.
The last sands grade into clays with oysters, beach and dune sands, overlain by
V-.!,1 biocalcarenites ("Calcarios com Chlamys scabrella de Casal Vistoso"). This
corresponds to a brief marine transgression (T3).
The next regression is marked by fluviatile V-.!,2. sands with Deinotherium. These
beds grade laterally into estuarine, more or less brackish facies, and these grade into
littoral marine calcarenites (V-.!_3, "Ca~carios com Chlamys scabriuscula de
Musgueira 0 ) (T4). V-~ sands directly overlie these beds.
Thus, transgressions TZ, T3, and T4 frame the Gomr,hotherium datum and
Deinotherium datum, respectively. Distribution of planktonicoraminifera, according
to M.L. Ubaldo and G. Bizon is as follows (table 1), including data for TS (next trans-
gression). Even if a reappraisal is needed, the correspondence to Blow's planktonic
zones cannot change very much.
On this evidence, unit II approximately corresponds to N 5; transgression TZ spans

from N6 to N7; IV-~ to N7; transgression T3 to N8 (in part); V-.!_2. to N8; transgression
T4 still to N8; V-b probably, by extrapolation, to N9; transgression TS to N9 at first,
yet subsequent beds attain N13 at least.
256
Table 1. *
Distribution of planktonic Foraminifera in the successive transgressive
events TZ to TS {lower levels) recognized in the Lisbon Miocene series.
TZ T3 T4 TS
Globigerinoides guadrilobatus altia:eerturus Bolli +

Globigerinoides guadrilobatus immaturus Le Roy + + + +
Globigerinoides guadrilobatus Eimordius Blow 8t Banner +
Globigerinoides guadrilobatus guadrilobatus (D'Orbigny) + + +
Globigerinoides guadrilobatus trilobus (Reuss) + + +
Globigerinoides sicanus De Stefani + +
Globigerinoides subguadratus Bronnimann + + +
Globigerina angustiumbilicata Bolli +
Globigerina cf. ci:eeroensis Bolli +
Globigerina :eraebulloides Blow + + + +
Globoguadrina dehiscens (Chapman, Parr 8t Collins) + +
Globoguadrina larmeui Akers +
Turborotalia acrostoma (Wezel) +
Turborotalia meyeri (Cushman 8t Ellisor) +
Turborotalia obesa (Bolli) + + + +
Turborotalia eri eroronda Blow 8t Banner +
Turborotalia siakensis Le Roy) +
Orbulina suturalis Bronnimann +
*No recent revision has been made.
Ostracoda (according to A. Nascimento) provide additional evidence. IV-b

yielded Miociforideis fortisensis and Neocyprideis aguitanica, from the upper part of
Miocyprideis ortisensis zone m the Tagus basin. This species is well known elsewhere
as in the Aquitaine basin, yet never after Burdigalian. In V--.!1 there are but unchar-
acteristic brackish associations, and none is known in V-!!_. Hence an upper limit for
the Deinotherium datum only can be drawn still higher in V-!:.1 whose ostracofauna is
clearly of middle Miocene type and closely similar to more or less contemporary ones
in Aquitaine; it is marked by the appearance of Olimfalunia costata. Somewhat later
at VI-.!_ ("Argilas axuis de Xabregas") there is also Nonurocythereis seminulum. Both
species are absent everywhere prior to middle Miocene.
Evidence from calcareous nannoplankton is compatible. Upper IV-_!, only a little

under IV-~ yielded an association that may be ascribed to NN4 Martini zone (Fonseca,
1977, p. 74). This is in good agreement with evidence based on planktonic foramini-
fera. No data are available for IV-b to V-b units.
All in all, the Deinotherium datum falls approximately in Blow's NS zone, or at

least the first part of it. The V-!!_ "His:eanotherium fauna" may still not be younger
than NS; stratigraphically it underlies V-_£ beds with Foraminifera indicating N9,
Globorotalia :eeriJ)heroronda zone, upper Langhian. This is reinforced by the presence
of Olimfalunia costata and calcareous nannoplankton (according to C. Muller) indi-
cating the middle Miocene NNS, S:ehenolithus heteromoryhus zone.
Pollen and spores do not add much to the age of the proboscidean data. How-
ever, they offer much valuable information about climate and ecology.
Age of Proboscideans Data Placed in the

Paleogeographical aDd Geotectonic Conte:d
There is a gap between the II and IV-b faunas, from about ZO to near 17 Ma.
Nonmarine mammals are unknown in m and IV-a faunas.
257
Mammalian faunas from upper I and n are different from the IV-.!!_ fauna as a
consequence of migration, local evolution, and maybe also some extinctions.
This situation differs from that concerning IV-]! and V-.!_ faunas, the last being
about 16.5 Ma in age. In spite of the short time span, there are faunal differences. In
situ evolution really occurred within some taxa but it is not enough to explain most
differences. These can instead be assigned to immigration of new taxa through a
pathway that probably did not work before. Arrivals of Deinotherium, Bunolistriodon,
and Gaindatherium (Iberotherium) may reflect this new pathway (figure Z).
The event that may have completed this pathway was the collision between
Betic and Hesperic massifs, thus closing a westward gap through the Alpine Arch into
Iberia (cf. Antunes, 1979). Mammal migration to Iberia may not have happened only
through the "normal" European way from France but (at least episodically) in the
reverse sense, as it seems evident after the rare but sure presence of both
Hispanotherium and G. (Iberotherium) in some French early middle Miocene sites
(Ginsburg et al., 1987).
late
AF
Fig. z. Proboscidean data: Expansion in early and early

middle Miocene. Thick line = first gomphothere
expansion (late early Miocene) from Africa into
Eurasia, the Indo-Pakistan subcontinent, Japan,
and North America (and much later to South
America), westernmost Europe being invaded
through the "normal" European way. Thin line=
first deinothere expansion (latest early Miocene)
from Africa into Indo-Pakistan subcontinent and
Europe but not into eastern Asia and America
through (if maybe not exclusively) the Alpine
Arch; deinotheres probably were unable to travel
through rather arid regions (heavy dots) that
acted as ecological, selective barriers. (Later
dispersal of Proboscideans not considered here.)
258
As stated by Tobien (1986, p. 168-169) about Deinotherium, its arrival in Europe
proceeded along a way comprising Arabia, Asia Minor, and the Iberian Peninsula. This
expansion was also responsible for the spreading of deinotheres into the Indo-Pakistan
subcontinent (ibid.).
Of course, the major eustatic transgressions just before (Burdigalian, ill and IV-a
Lisbon stratigraphical units) and after (Langhian and Serravallian, V-c and VI-a}
proboscidean dispersal exerted a strong influence. The two brief episodescorrespond-
ing to subunits V-.!_1 and V~3 are related to tectonic events of more limited, regional
scope, like the collision referred to above.
The paleogeographical situation was not the sole factor controlling the migration
of mammals. Another leading factor was climate.
PALEOECOLOGY,CLEMATE,PALEOGEOGRAPHICAL,
AND ECOLOGICAL CONTROL
The Tagus basin provided much data of paleoclimatological interest (Antunes and
Pais, 1983; Pais, 1986). What may be of interest to frame the proboscidean data will
be summarized as follows.
During Aquitanian (unit I) a limited marine transgression occurred (TO), soon

followed by another and more important one (Tl). Conditions allowed the develop-
ment of fringing reefs, which never could be built later. Bombax and other tropical or
subtropical trees and shrubs indicate the presence of lowland forests. Moist and warm
conditions must have prevailed.
As for the next regressive event, lagoons were well developed, soon to be super-
seded by swamps with brackish or fresh waters. Sands were deposited by rivers.
Upper unit I mammals (anthracothere, some rhinocerotids) indicate swamp and forest
environments. Not much later (unit II) climate became less humid and less warm;
small mammals indicate open spaces in a steppe-like environment with bush and
scarce trees (Antunes and Mein, 1986). There is a good fit between plant (Pais, 1986,
p. 189) and small mammal (Antunes and Mein, 1986, p. 133-134) data.
Climate changed during the following transgression, TZ (units ill and IV-.!), and
became somewhat warmer and more humid, probably tropical. Warm sea waters
brought stenotherm Foraminifera (miogypsinids, heterosteginids), pelecypods such as
Avicula, and nautiloids (Aturia), as well as fishes.
The following regression (RZ) allowed the first immigration of proboscideans

(Gomphotherium), now represented in IV-.!!_ fauna. As in similar situations, a decrease
in humidity occurred (Antunes and Pais, 1983, p. 86, fig. Z; Pais, 1986, fig. 4).
Evidence from the Lisbon region is not enough to be sure if Deinotherium already
existed elsewhere, but there is not the faintest trace of Deinotherium in the IV-b
fauna. It almost certainly did not accompany the first gomphotheres to this area. -
Climate and environment must have accounted for the filtering that resulted in
differential migration, since no credible physical barrier could simultaneously permit
the gomphothere migration and prevent that of deinotheres.
In a brief time span (in uppermost Burdigalian) two short-lived transgressive

events (T3 and T4) occurred (V-.!_1 and V-.!_3 units), bringing in tropical sea waters and
stenotherm, thermophilic forms such as nurse sharks, large barracudas, and others. A
temperature and humidity optimum corresponding to tropical conditions was
attained. This is confirmed by the presence of several crocodilians including a
tomistomine and a probable gavial. Forest would then be much more developed on
land.
259
It is in such context that the first Deinotherium immigration occurred. It
undoubtedly corresponds to the local development of forests.
A counter proof is given by the much drier and rather arid environments prevail-
ing in earlier middle Miocene. Deinotheres would be represented (V-b fauna) but by
one tooth of doubtful origin, that is nearly nothing against hundreds of gomphothere
remains. The near absence in the V-b fauna of Deinotherium can be ascribed to the
deterioration of climate and to forestregression; evidence, mainly concerning plants,
leaves no doubt about this. We may, therefore, conclude the deinotheres were much
more demanding than gomphotheres, and much more dependent on humid and perhaps
riparian forest environments (as for the vegetation characters, see Pais, 1986, p. 189,
fig. Z-4).
In short, we must take into account both physical/physiographical situations and

climate for correctly placing the two proboscidean dispersal events at least in
westernmost Eurasia.
Paleogeography is one of the leading factors controlling mammalian migration.

Much limited by its own requirements, Deinotherium was unable to travel across dry
regions. Probably as a consequence, its first immigration into western Europe may
have occurred through the Alpine Arch, that was complete by the end of early
Miocene (and lasted until sometime prior to the great upper Langhian-Serravallian
transgression, which may have disrupted that pathway). Even so, the same pathway
was also available for gomphotheres; indeed, they largely exceed deinotheres in
numbers everywhere, even when the last were more common, as in the V-~ fauna, and
they could have traveled together (beside those already existing there).
As the Alpine Arch was rather narrow and surrounded by the sea, climate would
have been much more humid than in the inner regions of Eurasia. Large continental
areas acted as selective barriers, allowing the migration of gomphotheres (if not the
colonization by them), but not that of the more strictly conditioned deinotheres.
These two immigration waves into western Europe must have undertaken differ-
ent directions. This would be consistent with the facts that Deinotherium (and
gomphotheres) could, and did, attain the Indian-Pakistan subcontinent but never
reached East Asia (Tobien, 1986, p. 169), which required long travel across arid
areas. Gomphotheres were able to make their way as far as Japan and further on into
North America. Deinotheres could not do likewise; control must have been of a
climatic-ecological nature.
We can thus summarize the succession of events as follows:
At about 17 Ma, in westermost Europe climate was subtropical with drier areas;
gomphotheres arrived for the first time through the "normal" European way (= the
Gomphotherium datum);
Approximately at 16 Ma the Alpine Arch closed, and presented an alternate path-

way; Iberian climate was probably then tropical with an optimum of warmth and
humidity; two regional tectonic events related to the collision between Hesperic
and Betic massifs are linked to two brief transgressive oscillations, framing a
regression episode marked by the arrival, probably through the Alpine Arch, of
deinotheres (= the Deinotherium datum).
Regression and probably a much drier climate - maybe in close relationship with
the restriction of Tethys and Paratethys seas -- drastically changed the environ-
ments, and became distinctly arid in westernmost Europe; a new migration wave
reached Iberia through the Alpine Arch (probably the shorter means of land
communication with the East) bringing in Hispanotherium, Chilotherium, and other
forms adapted to steppe, more or less arid environments; mammal migration then
proceeded eastward across the Pyrenees.
260
The great eustatic middle Miocene (Langhian-Serravallian) transgression inter-
rupted communications through the Alpine Arch, thus closing its role as a land
bridge.
CONCLUSIONS
1. Lower Tagus basin, being the westernmost Old World distribution area for Miocene
Proboscidea, is of particular interest to check all evidence about their first
arrivals.
z. Regional evidence allows a multidisciplinary approach giving an overall view about

age, correlation between marine and nonmarine faunal associations and events,
ecology, climate, and paleogeography.
3. There are indeed two events, a Gomphotherium datum and a Deinotherium datum.
4. Both can be framed chronologically: the Gomphotherium datum at about Blow's N7

zone, Burdigalian, approximately 17 ± 0.5 Ma; and the Deinotherium datum at N8,
uppermost Burdigalian, about 16 ± 0.5 Ma.
5. Gomphothere and deinothere migrations from their dispersal center in Africa were
brief events of probably not more than 1 Ma, even for the farthest extreme of
gomphothere distribution range in North America.
6. Migrations of proboscidea into westernmost Eurasia were acomplished through two

pathways, (a) across eastern Europe into the west until Iberia, as gomphotheres did,
and (b) also westward through the Alpine Arch - complete some 16.5 Ma ago -
into Iberia then eastward across the Pyrenees, this having been followed by
d~inotheres (and also by gomphotheres, too).
7. The main controlling migration factors were paleogeography and climate, arid
regions playing a filter role as they rejected migration of high-humidity demanding
forms as deinotheres, but not of savanna or steppe forms.
8. Deinotheres migrated into Iberia during a brief, warm and humid episode; the same
Alpine Arch passage was still in use in early middle Miocene by mammals adapted
to drier environments like Hispanotherium, which migrated into France like
Deinotherium but when climate was much drier.
9. Alpine Arch passage was disrupted by the great middle Miocene transgression and
ceased to exist.
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Mainzer Becken (BRD). Mainzer Naturw. Archiv, v. Z4, p. 155-Z61.
Tobien, H., 1987. Bemerkungen zur Altersstellung der altmioziinen Saugerfauna von
Frankfurt/Nordbassin und der prabasaltischen Sedimentfolgen im Untergrund von
Frankfurt am Main. Geol. Jb. Hessen, v. 115, p. Z05-Z16.
Torres, A.S., 1935. Mais um dente de Mastodon (Tetrabelodon) angustidens, Cuv.
colhido no terciario de Lisboa. Bol. do Mus. Lab. Miner. Geol. da Univ. de
Lisboa, za 11t!rie~ no. 4, P• 41-44.
Zbyszewski, G., 1937. J;)ecouverte de nouveaux gisements terrestres dans le Neogene
des environs de Lisbonne (Portugal). C. R. Acad. Sci. Paris, t. Z05, p. 1Z41-1Z4Z.
Zbyszewski, G., 1941. Note sur Ia decouverte du genre Dinotherium au Portugal.
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Zbyszewski, G., 1949. Les vertebres du Burdigalien superieur de Lisbonne. Serv. Geol.
de Portugal, Lisboa, 77 p.
262
PATTERNS OF OLD WORLD BIPPARIONINE EVOLUTIONARY
DIVERSIFICATION AND mOGEOGRAPWC EXTENSION
R.L.Bemor
College of Medicine
Department of Anatomy
Laboratory of Paleobiology
Howard University
Washington D.C. Z0059 and
Department of Paleobiology
Museum of Natural History
Smithsonian Institution
Washington D.C. Z0560
H. Tobien
Johannes Gutenberg Universit'at
Institut fUr Geowissenschaften FB ZZ
Saarstrasse Z1, Postfach 3980
D6500, Mainz, West Germany
M. 0. Woodburne
Department of Earth Sciences
University of California
Riverside, California 9Z5Z1
INTRODUCTION
Hipparionine horses have long been united evolutionarily by the presence of

three toes per digit, having high crowned cheek teeth with cement, and isolated proto-
cones on upper cheek teeth (Christal, 183Z). Geochronologically they have further
been recognized as the preeminent large mammal "index" fossils for late Neogene Old
World deposits. Their abundance in later Neogene mammal faunas has prompted the
production of a staggering body of systematic and interpretive literature during the
last 150 years. In the last 40 years there has been an increasing number of attempts
to reorganize parts of Old World hipparionine systematics by regional studies of vari-
able scope including Europe in general (Pirlot, 1956); western Europe (Rhone Valley:
Sondaar, 1974; Spain: Alberdi, 1974); eastern Europe, western U.S.S.R. and North Asia
(Gromova, 195Z; Gabunja, 1959; Zhegallo, 1971, 1978); Europe, North Africa, and
southwest Asia (Woodburne and Bernor, 1980); southwest Asia (Bernor, 1985a); China
(Qiu et al., 1987; Bernor et al., in press); the Indian Subcontinent (Hussain, 1971;
MacFadden and Woodburne, 198Z; Bernor and Hussain, 1985); and Africa (Bone and
Singer, 1965; Churcher and Richardson, 1978). Extensive interregional comparisons of
Old World hipparionines have been provided by Gromova (195Z) and ForstEm (1968).
The studies cited here have been preceded by a myriad of more isolated ones,
which can be extracted from the literature reviewed here. Recent summaries of Old

Edited by E. H. Lindsay et at.
World evolutionary relationships have been provided by Woodburne and Bernor (1980),
Bernor and Hussain (1985), Bernor et al. (1987b), Woodburne (in press), and Alberdi (in
press). A major review of phylogenetically relevant North American hipparionines has
been made by MacFadden (1984), while Hulbert and MacFadden (in review) have made
an important study of the phylogenetic relationships of parahippine and merychippine
sister taxa.
We review here several selected Old World hipparion species and potential North
American sister-taxa. Our selection is made to emphasize the recognition of discrete
evolutionary lineages which have emerged from work performed during the last 15
years. We give here updated information on the discrete morphological states (table
1) which distinguish these species, and group sets of them into lineages. Summary
statistics of cheek tooth row length and metapodial m maximum lengths and distal
articular widths are tabulated, when known (table 2). Tables 3 and 4 give bivariate
measurements for calcanea and astragali. We then review the various competing
hypotheses of these superspecific phylogenetic relationships, and the distribution of
the clades in time and space. Our essay serves to further suggest that Old World
hipparions are represented by several superspecific groups of at least genus-rank, and
that the group can be usefully applied for provincial to intercontinental-scale
correlations.
CONVENTIONS AND ABBREVIATIONS
Temporal Terms
These terms are specific to the continent in question, without repetition in the
text. Some geographical divisions of the Old World are recognized (in parentheses) to
aid in biogeographic discussions. New World ages follow Tedford et al. (1987), Old
World ages follow Berggren et al. (1985a,b), Qiu et al. (1987; specifically China),
except where otherwise cited. We emphasize that the ages of various equid species
referred to here have substantially varying degrees of chronologie resolution. Gener-
ally speaking, North American equids have the highest degree of resolution, based on
an extensive data matrix of biochronologic, isotopic, and magnetostratigraphic corre-
lations. Most European ages are based on mammal Neogene correlations (MN zones;
see Mein, this volume). Although an integrated mammalian biochronology/magneto-
chronology is now emerging, largely due to the efforts of Sen (see manuscript in this
volume), most European localities that produce hipparions are stratigraphically uncon-
fined and calibrated only indirectly. Furthermore, study of some Spanish units (with a
well-developed local stratigraphy) has led Daams and Freudenthal (1981) and Daams et
al. (1987) to suggest that at least some of the MN zonations have limited geographic
utility. For all of these reasons, assignment of isotopic age correlations made with
respect to European sites (and thus hipparion species) should be taken cautiously.
Indo-Pakistan hipparion records are similarly uncertain because of the poorly known
provenance for a large proportion of the better fossil material and relatively few
calibration points, either directly or by magnetostratigraphic correlation. China's
recQord is improving due to the efforts by the Institute of Vertebrate Paleontology and
Paleoanthropology to strike both biochronologic correlations with the European
mammal biochronology framework, as well as to develop an independent isotopic and
magnetostratigraphic chronology (see Qiu, this volume). At the moment, however,
this work is in early stages, relying mostly on biochronologic correlations to the
European MN zonation, and an independent calibration network has not yet been
developed. East Africa has a mature isotopic chronology which provides a number of
important tie-points for provincial correlations between East African and other Old
World faunas. For the bulk of the European and Asian hipparion species, assignment of
isotopic ages should be taken cautiously, however.
264
Table 1. Character state distribution on selected Old World hipparionines and their related sister-taxa.
Taxa 2 3 4 5 6 7 8 9 10 ll 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40
P. leonensis A A A D C A B/C B A A A A A A A A A C A D B D A A A B
NA, 19-18 Ma
2 !!_· ins ignis A A A C B A B A A A A A A A A A A B/C A B A/B D B A/B B A
NA, 16-15 Ma
3 .£• goorisi B A C A A A B A B B A A A A A ?A .B B/C A B B D B B ?B A A A A A A A A A B B B B A A
NA, 15 Ma
4 .£• sphenodus B A B/C A A A A A A/B B A A A A/B A B B/C B/C B B B A B C B B A A A A A A A B B A B A A
NA, 14-12 Ma
5 c. occidentale C ?B B A A A A A B B A A A C B B C B B A/B B A B C B A A A A A A A A A B B B B A A/C
NA, 12-8 Ma
6 .£• plicatile ?C ?B A B A A B C
NA, 9-6 Ma
7 .£• ingenuum ?C ?B A B A A B C C B B c
NA, ll-6 Ma
8 C. emsliei ?C ?B A A B C+ A B A B/C B B C B A AAAAAA B BB A A
NA, 5-2 Ma
9 ':!!.•" shirley! B ?A C B B A B B A B A A A C A B A C A B/D A C B B B B A A A A A A A B B c B B A A
NA, 14 Ma
l 0 "H." tehonense c C B B A B B B B A A A C A B C C A A/B C C B C B B A A A A A A A A B C B A A B
NA, 12-9 Ma
ll "H." force! c C/K C B A C B B B A A A C B B C B/C A A/B C C B C B B A A A A A B B B B A A/B
NA, ll-6 Ma
12 ':!!_." primigenium C B B A A A A A B B A A A C B B C A B A B B B C B A A A A A A A A A B A/B A B A B
~·~· Eu, ll-8 Ma
13 "H." catalaunicum C B H A A A A A B B A A A C B B C A B A B B B B B A
E~, ll Ma
14 "H. mediterraneum" C B I D CACBBBAAA CBB C BB A B DB C B A
E~, ca. 8 Ma
15':!!.·"~ CB C A BABBBBAAA CBB C BB A C C B C B A
SWA, 9 Ma
16 "H." melendezi C B J A B A B B B B A A A C B B C A B A B/C B B C B A
N E~, 10 Ma
C»
(11 (cont.)
N
en
en Table 1. (continued)
Taxa 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40
17 ·~·" giganteum CB B B BAABBBAAA CBB CAB A C BB CBA A A A A

WAs, 8 Ma
18 "H." africanum C B H A A A A A B B A A A D B B C A B A B A/B B C B A A A A A A A A A B A/B A B B/C B/C
NAf, 10.5-5 Ma
19 "H." weihoense CB C A AAAABBAAA CBB CAB A B AB CBA
EA, 11-10 Ma
20 "H." dermatorhinum D B B B B A A A B B A A A F B B C A B A/B B G B C B A A A A B B B B A B B/C A/B B A/B C
EA, 10-7.5 Ma
21 ·~·" coelophyes C B D C B/C A B/C B B B A A A C B B C A B A B B B C B A
EA, 8 Ma
22 '~·" hippidiodus E B G D C A D B B B A A A B B C B/C B B C/D C/D B C B A
EA, 8-6 Ma
23 prostylum
.!!.· D B D B/C B A B/C B B B A A A C B B C B B A/B B/C C B C B A A A A A A A A A B B B B B/C B/C
Eu, SWA, 8.5 Ma
24 H. dietrichi D B F C/D C A D B B B A A A C B B C B B B C C B C B A
SWA, 8-7 Ma
25 .!!.· campbelli D B F C/D C A D B B B A A A D B B C B B B C C B C B A A A A A A A A A B C B B A C
SWA, 7 Ma
26 .!!.· antelopinum ?D B ?C ?C C A ?C B B B A A A B B C B B B C C B C B A
SA, L.M.
27 ':2.·" platyodus C B B B/C B A B A B B A A A C B B C A B A B B B C B A A A A C C C C A B B/C A B A D

EA, 9. 5-5 Ma
28 ':2.·" ptychodus G B J B A A B A B B A A A ?C B B C A+/A B A/C B B B C B A
EA, 8-4.5 Ma
29 ':2.·" perimense GB DC BAAABBAAA CBB DAB A B EB CBAAAACCCCAB A CB D D
SA, 8 Ma
30 "S." houfenense EB F D DADBBBAAA CBB DAB A B EB CBABAACCCCAB A AB D D
EA, 6-2.5 Ma
31 "S." aff. HB G E DAEBBBAAA CBB D BB A B EB CBABAACCCCAB B B D D
houfenense
EAs, ca. 2 Ma
Taxa 2 3 4 5 6 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40
32 "S." r. crusafonti H B G E D A E B B B A A A BB DAB A B EB CBABAACCCCAB B D D

E~, ca. 3 Ma
33 "S." turkanense ?GB G D CADBBBAAA CBB DAB A B EB CBA
if., 5-4 Ma
34 "S." af arense ?G B G C C A D B B B ?A A A ?C B B D A B A B C/E B C B A B B B C C C C A B A/B B A D D
EA, 3 Ma
pater H B G E D A E B B B A A A G B B D A+ B A B B B C B A A A A C C C C A B B D D
35 .!.·
EA, 5-2 Ma
36 P. sinense H B G E D A E B B B A A A H B B D A+ B C B F B C B A A A A C C C C A B C B B D D
EA, 3-2 Ma
37 C. moldavicum B B C B A A A A B B A A A C B B C B B A/B C D B C B A A A A A A A A A B C B B A/B B
SWA, 9-8 Ma
38 C. matthew! B B B C C A B B B B A A A C B B C B B A/B C B/D B C B B A A A A A A A A B C A B A C
SWA, 8.5-7 Ma
39 C. mediterrsneum B B B B A A A A B B A B A D B B C A B A B B/D B C A A
SWA, 8 Ma
40 .£• proboscideum B B B B A A A A B B A B A E B B C A B A B B/D B C B A
SWA, 8-7 Ma
41 C. aff. B B B B A A A A B B A B A D B B C A B A B B/D B C B A
-;;;'editerraneum
EA, L.M.
42 c. forstenae B B B C C A C B B B A B A E B B C B B A/B C/D C/D B C B A
EA, 9-7.5 Ma
43 C licenti BB B B ABAABBBBB GBB C CB B D DB CBA
EA, 5-3 Ma
(continued)
...,
0)
......
Table 1. (continued) Legend
~·leonensis (= Parahippus leonensis); references= Hulbert and MacFadden (in review)

M. insignis (= Merychippus insignis); references = Skinner and Taylor (1967;
measurements); Hulbert and MacFadden (in review)
C. goorisi (= Cormohipparion goorisi); references= Woodburne et al. (1981);
MacFadden (1984; measurements) ; Hulbert and MacFadden (in review)
C. sphenodus (= Cormohipparion sphenodus); Woodbume et al. (1981); MacFadden
(1984; measurements)
C. occidentale (= Cormohipparion occidentale); Woodburne et al. (1981}; MacFadden
(1984; measurements); Bernor and Hussain (1985); Bernor et al. (1987); Bemor et
al. (1988)
C. plicatale (= Cormohipparion plicatile); MacFadden (1984); Hulbert (in press)
C. ingenuum (- Cormohipparion ingenuum); MacFadden (1984); Hulbert (in press)
C. emsliei (= Cormohipparion emsliei); Hulbert (1987) (PZ/-M3/ measurement
estimated here)
"H." shirleyi (= "Hipparion" shirleyi); Woodburne et al. (1981); MacFadden (1984;
measurements)
"H." tehonense (= "Hipparion" tehonense); MacFadden (1984; measurements)
"H." forcei (= "Hipparion" forcei); MacFadden (1984; measurements)
"H." primigenium s.s. (= "Hipparion" primigenium s.~.); Tobien (1986); Bernor and
Hussain (1985); Bernor et al. (1987b); Bernor et al. (1988); Bernor et al. (in press;
measurements)
"H." catalaunicum (= "Hipparion" catalaunicum); Woodburne and Bernor (1980); Bernor
(1985); Bernor et al. (1987b); measurements from Bernor (personal observation)
"H. mediterraneum " (= "Hipparion mediterraneum " sensu Alberdi, 1974; sic); Alberdi
(1974); measurements from Bernor (personal observation)
"H."~(= "Hipparion" ~; Woodburne and Bemor (1980); Bernor (1985a);
Bernor et al. (1987b)
"H." melendezi (= "Hipparion" melendezi); Alberdi (1974; measurements); Woodburne
and Bernor (1980); Bernor and Hussain (1985); Bernor et al. (1987b)
"H." giganteum (= "Hipparion" giganteum); Gromova (1952); Woodburne and Bernor
(1980); Bernor et al. (1987b); measurement from Bernor (personal observation)
"H." africanum (= "Hipparion" africanum); Arambourg (1959); Woodburne and Bernor
(1980); Bernor et al. (1987a; measurements); Bernor et al. (1987b); Bernor et al.
(1988)
"H." weihoense (= "Hipparion" weihoense); Liu et al. (1978); Bernor et al. (1988);
measurements from Bemor (personal observation)
"H." dermatorhinum (= "Hipparion" dermatorhinum ); Bemor et al. (1987b); Qiu et al.
(1988; measurements); Bernor et al. (1988); Bernor et al. (in press)
"H." coelophyes (= "Hipparion" coelophyes); Bernor et al. (1987b); Qiu et al. (1987);
Bernor et al. (1988); Bernor et al. (in press; measurements)
"H." hippidiodus (= "Hipparion" hippidiodus); Bernor et al. (1987b); Qiu et al. (1987);
Bernor et al. (in press; measurements)
H. prostylum (= Hipparion prostylum); Woodburne and Bernor (1980; measurements) ;
Bernor et al. (1980); Bernor (1985a; measurements); Bernor et al. (1987b)
H. dietrichi (= Hipparion dietrichi); Woodburne and Bernor (1980); Bernor et al. (1980);
Bemor (1985a); measurements from Bernor (personal observation)
H. campbelli (= Hipparion campbelli); Bernor (1985ar
H. antelopinum (= Hipparion antelopinum); Woodbume and Bernor (1980); MacFadden
and Woodburne (1982); Bernor and Hussain (1985); measurements from Bernor
(personal observation)
".§_." platyodus (= "Sivalhippus" platyodus); Bernor et al. (1987b); Qiu et al. (1987);
".§_." ptychodus (= "Sivalhippus" ptychodus); Bernor et al. (1987b); Qiu et al. (1987);
"~." perimense (= "Sivalhippus" perimense); MacFadden and Woodburne (1982); Bemor
and Hussain (1985; measurements)
"~." houfenense (= "Sivalhippus" houfenense); Flynn and Bernor (1987); Qiu et al. (1987)
"~." aff. houfenense (= "Sivalhippus" aff. houfenense); MacFadden (1984}; Flynn and
Bernor (1987)
268
"~." !.• crusafonti (= "Sivalhippu s" rocinantis crusafonti ); Alberdi (1974; cheek tooth and
metapodia l distal articular measurem ents estimated from figures)
"~." turkanense (= "Sivalhippu s" turkanense ); Hooijer and Maglio (1976; measurem
ents);
Bernor and Hussain (1985)
·~." afarense (= "Siva1hipp us" afarense); Eisenmann (1976; measurem ents)
_E. pater (= Proboscidi pparion pater); Qiu et al. (1988; measurem ents)
P. sinense (= Proboscidi pparion sinense); Qiu et al. (1988; postcrania l measurem ents);
Bernor et al. Un press; skull measurem ents)
C. moldavicu m (= Cremohipp arion moldavicu m); Gromova (1952); Bernor (1985a;
measurem ents)
C. matthewi (= Cremohipp arion matthewi) ; Bernor (1985a); measurem ents from Bernor
(personal observatio n)
C. mediterran eum (= Cremohipp arion mediterran eum); Woodburne and Bernor (1980);
Bernor (1985a); measurem ents from Bernor (personal observatio n)
C. proboscide um (= Cremohipp arion proboscide um); Sondaar (1974); Woodburne and
Bernor (1980); Bernor (1985a); measurem ents from Bernor (personal observatio n)
C. aff. mediterran eum (= Cremohipp arion aff. mediterran eum); Bernor (personal
observatio n)
C. forstenae (= Cremohipp arion forstenae) ; Bernor et al. (1987b); Qiu et al. (1987);
Bernor et al. (in press; measurem ents)
C. licenti (= Cremohi arion licenti); Bernor et al. (1987b); Qiu et al. (1988;
- -measure ments ; Bernor et al. (in press)
NA = North America
Eu =Europe
SWA =Southwe st Asia (includes Greek and western U.S.S.R. localities)
NAf =North Africa
EAf = East Africa
EA = East Asia
SA = South Asia
L.M. = late Miocene
269
...,
..... Table 2.. Measurements on selected hipparionine maxillary cheek tooth rows, Me m•s and Mt ID's from Holarctica and Ethiopia
0
Mcm Mtm
Length P2./-M3/ Length DAW Length DAW
Taxon g OR g OR g OR g OR g OR
1 P. leonensis
NA, 19-18 Ma
2. M. insignis 116.4 108.9-12.1.4
NA, 16-15 Ma (n=7)
3 C. goorisi 116.8 111.2.-12.1.3
NA, 15 Ma (n=4)
4 C. sphenodus 12.6.5 12.0.2.-132..8
NA, 14-12. Ma (n=9)
5 C. occidentale 138.0 12.5.1-151.0
NA, 12.-8 Ma (n=13)
6 C. plicatile 12.5.5
NA, 9-6 Ma (n=Z)
7 C. ingenuum 115.9
NA, 11-6 Ma (n=1)
8 C. emsliei 135.0 2.34.0 35.5
NA, 5-2. Ma (n=l) (n=1) (n=1)
9 "H." shirleyi 105.4 97.8-106.6 147.6 147.2.-148.0 2.0.9 2.0 .2.-2.1.6 166.3 165.8-167.8 18.5 18.2.-18.8
NA, 14 Ma (n=4)
10 "H." tehonense 12.3.9 110.2.-134.6
NA, 12.-9 Ma (n=36)
11 "H." forcei 133.5 132..7-134.3
NA, 11-6 Ma (n=2.)
12.a "H." primigenium s.s. 2.15.6 37.5 2.40.2. 2.3 6.-2.4 7.3 38.0 36.2.-39.0
Eppelsheim, ca. 11 Ma (n=l) (n=1) (n=4) (n=4)
Mcm Mtm
Length PZ/-M3/ Length DAW Length DAW
Taxon :X OR :X OR :X OR :X OR :X OR
1Zb "H. n primigenium s.s. 158.7 147.0-167.8 Z1Z.7 Z03.0-ZZO.Z 36.7 33.7-39.9 Z4Z.6 ZZ3.9-Z5Z.3 37.6 34.9-41.1
Howenegg, ca. 11 Ma (n=7) (n=Z1) (n=Z1) (n=Z4) (n=Z4)
1Zc "H." primigenium s.s. 158.0 Z09.Z Z06.7-Z11.7 33.9 3Z.6-35.1 Z48.7 Z45.9-Z51.7 36.3 34.0-38.1
Charmoille, ca. 11 Ma (n=1) (n=Z) (n=3) (n=4) (n=5)
1Zd "H." primigenium s.s. Z07.8 Z01.8-Z14.0 37.0 33.9-39.1 Z31.7 ZZ4.5-Z38.5 35.3 34.4-36.5
Vienna basin, ca. 11 Ma (n=4) (n=4) (n=6) (n=6)
13 "H." catalaunicum 151.4 Z06.0 36.0 Z36.Z ZZ6.0-Z59.0 37.9 33.4-4Z.O
Eu, 11 Ma (n=1) (n=1) (n=1) (n=5) (n=4)
14 "H. mediterraneum"
Eu, ca. 8 Ma
15 gettyi
n H. n 137.8
SWA, 9 Ma (n=1)
16 "H." melendezi 143.1 Z14.5 Z14.0-Z15.0 31.3 Z9.5-33.Z ZZZ.5 Z18.5-ZZ5.7 33.3 Z9.6-37 .9
Eu, 10 Ma (n=1) (n=Z) (n=Z) (n=3) (n=3)
17 "H. n giganteum 154.7 146.8-16Z.8 Z11.3 197 .O-Z30 .0 37.9 36.0-41.6 Z44.9 ZZ4.0-Z6Z.O 38.9 35.0-4Z.O
WAs, 8 Ma (n=4) (n=67) (n=6Z) (n=76) (n=69)
18 "H." africanum 145.6 139.6-153.5 Z07.7 Z03.3-Z11.1 36.0 35.6-36.3 Z40.4 Z33.8-Z45.8 35.0 33.9-36.Z
NAf, 10.5-5 Ma (n=6) (n=3) (n=5) (n=5) (n=5)
19 w eilioense
n H." 168.5
EA, 11-10 Ma (n=l)
ZO n H." derma torhinum 151.Z 140.0-16Z.3
EA, 10-7.5 Ma
Z1 "H." coelophyes 131.0 1Z5.0-140.7
N
..... EA, 8 Ma (n=8)
(continued)
"'..... Table z. (continued)
"'
Mcm Mtm
Taxon X OR X OR X OR X OR X OR
ZZ "H." hippidodus 139.0 134.3-147.9 Z16.3 Z14.0-ZZO.O 3Z.7 3Z.0-33.4 Z44.0 31.Z
EA, 8-6 Ma (n=S) (n=4) (n=4) (n=1) (n=1)
Z3 H. prostylum 140.0 1ZZ.7-15Z.6 Z05.6 Z01.Z-Z17 .4 30.3 Z9.1-31.9 Z4Z.4 ZZ8.9-Z51.0 30.1 Z7.4-33.8
Eu, SW A, 8.5 Ma (n=9) (n=6) (n=S) (n=13) (n=1Z)
Z4 H. dietrichi 141.1 1Z1.Z-154.5 Z35.0 ZZ7 .O-Z46.0 35.9 34.5-38.4 Z81.0 Z73.0-Z88.0 30.1 Z8.9-31.3
SWA, 8-7 Ma (n=1Z) (n=8) (n=8) (n=8) (n=Z)
ZS H. campbelli 154.3 Z53.4 ZSZ.7-Z54.1 3Z.6 31.6-31.8 33.4
SWA, 7 Ma (n=1) (n=Z) (n=4) (n=1)
Z6 H. antelopinum 133.8
SA, L.M. (n=1)
Z7 "~." platyodus 141.1 131.1-149 17Z.O 36.0
EA, 9.5-5 Ma (n=4) (n=1) (n=1)
Z8 "~·" ptychodus 137.9 1Z8.4-147.4 17Z.O 35.9
EA, 8-4.5 Ma (n=3) (n=1) (n=1)
Z9 "~."perimense 150.6 133.3-17 5.5
SA, 8 Ma (n=S)
30 "S." boufenense 144.9 139.7-148.0 Z49.8 Z41.0-Z59.7 43.9 41.4-47.Z Z73.8 Z57 .Z-Z86.8 4Z.9 40.3-44.7
EA, 6-Z.S Ma (n=3) (n=ZO) (n=ZO) (n=Z1) (n=Z1)
31 "S." aff. houfenense
c"'i. Z Ma
3Z "S." r. crusafonti 160 Z41.7 Z35-ZSO 40.0 Z74.7 Z64-Z85 43
ca. 3 Ma (n=1) (n=1) (n=9) (n=1)
33 "S." turkanense 163.0 41.0
EAf, 5-4 Ma (n=l) (n=1)
Mcm Mtm
Taxon X OR X OR X OR X OR X OR
34 "S." afarense 164.0

EAf, 3 Ma (n=1)
35 P. pater 155.5 155.2..155.8 2.34.3 2.19.0-2.47.0 38.8 35.7-4Z.4 2.66.1 Z5Z.Q-Z87 .4 37.8 33.Z-4Z.O
EA, 5-Z Ma (n=Z) (n=8) (n=8) (n=9) (n=8)
36 P. sinense 164.5 Z74.5 Z58.Q-Z98.0 42..4 38.2.-47.1 32.0.3 2.98.6-350.0 42..5 37.3-47.7
EA, 3-Z Ma (n=1) (n=16) (n=15) (n=16) (n=17)
37 C. moldavicum 130.6 12.0.0-141.3 2.05.9 195.0-2.13.0 31.1 Z9.5-3Z.5 2.35.0 zz5.o-Z59.0 31.4 2.9.5-33.0
SWA, 9-8 Ma (n=7)
38 C. matthewi 116.9 111.8-12.1.9 Z08.4 Z00.1-ZZ8.3 Z6.6 Z3.5-Z8.8 ZZ6.6 2.13.7-2.39.4 2.5.1 Z4.Z-Z5.9
SW A, 8.5-7 Ma (n=Z) (n=7) (n=7) (n=Z) (n=Z)
39 C. mediterraneum 140.4 132..0-150.3 Z16.0 ZOZ.0-2.34.0 3Z.Z 2.9.5..,.35.3 2.37.3 2.10.0-2.70.0 3Z.4 Z9.3-36.6
SWA, 8 Ma (n=3) (n=39) (n=38) (n=61) (n=54)
40 C. proboscideum 149.0 135.5-162..1 Z44.5 2.43.0-2.46.0 34.9 34.0-39.0 2.73.6 2.68.0-2.78.5 35.4 35.0-36.0
SWA, 8-7 Ma (n=8) (n=4) (n=4) (n=5) (n=S)
41 C. aff. mediterraneum 131.8 1Z8.6-134.9
EA, L.M. (n=Z)
4Z C. forstenae 12.9.3 12.1.7-137.0
EA, 9-7.5 Ma (n=8)
43 C. licenti 12.9.4 12.4.5-134.3
EA, 5-3 Ma (n=Z)
Legend
Taxa - as listed in Table 1 X- mean
Length PZ/-M3/ - length of cheek tooth row P2./-M3/ at occlusal surface OR- observed range
Me m - Metacarpal m Mt m - Metatarsal m
...., Length - maximum length (measurement 1 of AMNH conference) Length, DAW, X, and OR
..... DAW- distal articular width (measurement 11 at AMNH conference) - as cited above for Me m
w
Temporal Characteristics and Defiuitious
Hemingfordian - mammal age utilized as an interval corresponding to part of early

Miocene time in North America, ca. ZQ-16 Ma.
Barstovian - mammal age utilized as an interval corresponding approximately to the

middle Miocene in North America, ca. 16-11.5 Ma.
Clarendonian - mammal age utilized as an interval corresponding to part of the late

Miocene in North America, ca. 11.5-9 Ma.
Hemphillian - mammal age utilized as an interval corresponding to part of the late

Miocene in North America, ca. 9-5 Ma.
Blancan - mammal age utilized as an interval corresponding to part of the Pliocene in

North America, ca. 5-1.9 Ma.
Vallesian - mammal age utilized as an interval corresponding to part of the late

Miocene time in the Old World, ca. 11.5-9.5 Ma (modifications after Bernor et
al., 1988). Furthermore, note that Berggren et al. (1985a, Fig. 6; 1985b, Fig. Z)
have revised the age of the early part of Chron C5N to be about 1Q-9.5 Ma.
Berggren et al. (1985b, Fig. 3) indicate, however, that the Vallesian/Turolian
boundary is calibrated at 9.5 Ma, and Barry et al. (1982., 1985) indicate that the
age of the part of Chron C5N that corresponds to the local FAD of "Hipparion"
is circa 9.5 Ma. For the purposes of this manuscript we follow Woodburne (in
press) in taking the Vallesian/Turolian boundary as ca. 9.5 Ma, and follow Barry
et al. (1982., 1985) in recognizing the "Hipparion" FAD in the Siwaliks as
9.5 Ma. Note that Badgley et al.'s report of a "Hipparion" FAD in the Siwaliks
was restricted to a single section, with a precisely developed sedimentary
record, and is not meant to represent the Siwalik provincial "Hipparion Datum"
(Barry, personal communication).
Turolian - mammal age utilized as an interval corresponding to the late Miocene in

Europe and West Asia, ca. 9.5-4.9 Ma.
Ruscinian - mammal age utilized as an interval corresponding to early Pliocene time

in Europe and West Asia, ca. 4.9-3.4 Ma.
Villafranchian - mammal age utilized as an interval corresponding to the later Plio-

cene in Europe and West Asia, ca. 3.4-1.6 Ma.
Bahean - Chinese mammal age utilized as an interval roughly corresponding to the

Vallesian of Europe.
Baodean - Chinese mammal age utilized as an interval roughly corresponding to the

Turolian of Europe.
Gaozhuangian - Chinese mammal age utilized as an interval roughly corresponding to

the early Ruscinian of Europe.
Youhean- Chinese mammal age utilized as an interval roughly corresponding to the

late Ruscinian and early Villafranchian of Europe.
Nihewanian - Chinese mammal age utilized as an interval roughly corresponding to the

medial to late Villafranchian of Europe.
Ma- Megannum in the isotopic time scale. Where necessary, all isotopic calibrations
have been recalculated according to the decay constants utilized by Steiger and
Jaeger (1977).
E. and L. - Early and Late, as subdivisions of temporal intervals, and in contrast to the
stratigraphic intervals Lower and Upper; e.g., Land U.
274
Morpl!olosical Terms
All morphological characteristics are made on adult individuals. For cheek tooth
states, citation of the various morphological states is confined to approximate middle
1/3 stage-of-wear, whenever possible. Problems arise in definition of several cheek
tooth character states. Woodburne (in press) has cited specific ranges of maxillary
cheek tooth plications in his scoring of these as being complex, moderate, or simple.
The reader should recognize that throughout the ontogeny of a single individual there
may be considerable variability in the actual number of plications. It is not surprising
then that within a single population of individuals there is considerable variability in
plication frequency. In most cases, complexly plicated bands have a deeper amplitude
in middle stage of wear. Scoring of this "character" is somewhat complicated. Scores
b - moderate complexity and c - simple complexity are considered to be not only a
quantitative loss of the number of plis per fossette, but a decreasing gradient of
depth. Numbers of pli caballins, degree of hypoglyph incision, protocone shape, and
isolation of protocone, likewise can be quite variable both ontogenetically within a
single individual, and between individuals, and must be judiciously evaluated at medial
1/3 stage-of-wear. We adhere to this principle here.
All measurements are in millimeters= mm.
Statistics for figures 4, 9, and 10 were calculated using the software package
STATGRAPffiCS at the Landessammlungen flir Naturkunde, Karlsruhe.
Systematic Characteristics and Definitions
hipparionine or hipparion - any horse with an isolated protocone of the maxillary

premolar and molar teeth and, as far as known, tridactyl feet, including species
of the following genera: Hipparion, Neohipparion, Nannippus, Cormohipparion,
Hippotherium, Proboscidipparion, "Sivalhippus", Cremohipparion, Pseudo-
hipparion, and Stylohipparion. Characteristics of these taxa can be found in
MacFadden (1984), Bernor and Hussain (1985), Webb and Hulbert (1986), Hulbert
(1987, 1988), and Woodburne (in press). We present here a number of competing
hypotheses of several of these species' phyletic relationships and leave future
characterizations to subsequent systematic studies.
"Hipparion"- hipparionine horses that belong to different lineages than those listed
above, or which cannot be readily placed within a particular lineage.
"Group 1" hipparions - demonstrated by Bernor et al. (1988) to be a paraphyletic group

of primitive Old World hipparionines. These are characterized as being medium
to large sized hipparions almost always with a long preorbital bar; lacrimal
clearly placed well posterior to the posterior rim of the preorbital fossa (POF);
there is no lacrimal foramen; POF is primitively subtriangular shaped and
anteroventrally oriented, but may transform in more derived species to being
elongate to egg-shaped or a diminished rounded structure in more derived taxa;
POF is most often anteroventrally, but in some species anteroposteriorly
oriented; posteriorly, POF is most commonly deeply pocketed, being greater
than 15 mm in its deepest place, but derived members may reduce depth signifi-
cantly; POF's medial depth is usually great, being more than 15 mm in its
deepest place, its medial wall never contains internal pits, it most usually has a
distinctly defined anterior rim and infraorbital foramen is placed inferior to, or
encroaches upon, the anteroventral border of the preorbital fossa; buccinator
fossa is distinct and never posteriorly pocketed, there is no depression for the
caninus muscle and there is never a malar fossa; nasal notch is usually retracted
midway between canine and anterior limit of PZ/, although in some derived
species it becomes retracted as far as P4/; when present, P1/ is usually very
reduced or absent, and clearly non-functional; maxillary cheek teeth are
moderately curved to straight, maximum crown height is between 40 and 60 mm;
enamel plications of the pre- and postfossettes are most commonly complex in
middle stage-of-wear; posterior wall of the postfossette is distinct; pli caballins
are nearly always double; hypoglyphs are nearly always deeply incised; protocone
275
varies from being flattened lingually and rounded labially to being more oval to
round-shaped, always separate from the protoloph, and rarely has a protoconal
spur (usually on PZ/ only, when present at all) that is aligned lingually with
respect to the hypocone; PZ anterostyle/paraconid (e.g., MacFadden, 1984, Fig.
4) is always elongate; mandibular incisors are never grooved, always curved and
I/3's are never reduced in size; all mandibular cheek teeth usually have rounded
metaconids and me tasty lids with the notable exception of •H. n dermatorhinum
which has elongated ones; premolar ectoflexids do not usually separate meta-
conid and metastylid, while they do so in the molars; a single to complex enamel
folding of the pli caballinid is common, and metaflexid-entoflexid borders are
often complexly folded, especially in more primitive members of the "group;•
protostylids may or may not be present on all mandibular cheek teeth except
P/Z; ectostylids are not known to occur in the group; linguaflexids vary in this
"group• being shallow, U-shaped and shallow V-shaped; metapodials are robust to
more slenderly built.
Hipparion !.-!.- - These are, generally speaking, medium sized hipparions. They char-
acteristically have a preorbital bar somewhat shortened compared to •aroup t•
horses, but with the same relationships of the lacrimal-posterior to the posterior
rim of the POF; there is no lacrimal foramen; the preorbital fossa is reduced
compared to "Group 1 n horses, and transforms between species from being egg-
shaped to C-shaped and finally vestigial; usually, when present, POF is antero-
posteriorly oriented; posterior pocketing is moderate to absent; .medial depth is
moderate to very shallow and lacks internal pits; POF peripheral border is
moderately expressed to lacking and anterior rim is absent; infraorbital foramen
encroaches upon the anteroventral border of the POF; the buccinator fossa is
distinct and separate from the area of the caninus fossa and unpocketed posteri-
orly; there is neither a caninus nor a malar fossa; nasal notch is consistently just
anterior to or above PZ/; P1/ is usually not present; maxillary cheek teeth are
generally straight-walled, have a maximum height between 40 and 60 mm, their
fossette borders are generally thinly banded and have moderately complex
enamel ornamentation; posterior wall of the postfossette is always distinct; pli
caballins vary from being double to single in some species, are uniformly single
in others; hypoglyphs are generally moderately deeply incised; protocones are
generally oval to rounded-shaped structures, they are always isolated from
protoloph until very late stage-of-wear, and are more lingually placed than the
hypocone; protoconal spurs are very rare to absent; PZ anterostyle/paraconid are
always elongate; mandibular incisors are not grooved, they are curved and I/3's
are not reduced in size; premolar and molar metaconids and metastylids are
usually rounded; premolar ectoflexids do not separate metaconid and metastylid,
while they do so in the molars; pli caballinids and ectostylids are generally
absent; protostylids are only rarely present; ectostylids are absent; linguaflexids
are generally shallow and V- to U-shaped on the cheek teeth; metapodials are
known to be relatively elongate and gracile.
"Sivalhippus Complex•- This complex may include more than a single genus. Horses
of this group are characterized as having a tendency to become very large sized;
they usually have a lacrimal far posterior to the posterior rim of the POF, when
POF is present; when present, the preorbital bar is long; POF has a variety of
shapes and may be anteroventrally or anteroposteriorly oriented; there is no
lacrimal foramen; posterior pocketing of POF is slight to absent, medial depth
varies from deep to shallow or absent, there are no internal pits; peripheral
outline and presence of anterior rim varies in strength between taxa depending
on presence or absence of POF; infraorbital foramen is placed inferior to, or
encroaching upon anteroventral border of POF when POF is present; buccinator
and canine fossae are never confluent, buccinator fossa is never pocketed poste-
riorly, caninus and malar fossae are never present; nasal notch is highly variable
in its position between species of this clade, being retracted to the level of PZ/
in some, while being retracted in others as far as the orbits; P1 is nonpersistent
to absent; the maxillary cheek teeth are straight, tend to be very high crowned
276
(most species in excess of 60 mm maximum crown height}; occlusal morphology
is much like "Group 1" horses, having complex plications of the pre- and post-
fossettes, double or complex pli caballins and deeply incised hypoglyphs;
posterior wall of postfossettes are always distinct; protocones typically have a
triangular-shaped to triangular-elongate morphology, are always isolated from
the protoloph, rarely, if ever, have a protoconal spur and are aligned lingually
with respect to the hypocone; PZ anterostyle/paraconid are elongate; mandibular
incisors tend to become grooved, and in the most advanced African species there
is the transformation from curved to straight and hypertrophied incisor morphol-
ogy with concommitant atrophy of I/3; lower cheek teeth evolve strongly-
angular-facing borders of metaconid and metastylid; ectoflexid does not
separate metaconid/metastylid in premolars, while doing so in the molars; pli
caballinids vary from being complex to absent; protostylids are commonly
present and may be strongly developed as columnar structures; ectostylids
become prominent and permanent features in more derived African species of
this group; linguaflexids are usually deep and U-shaped to accommodate occlu-
sion with the large protocones; metapodials tend to be elongate and quite robust-
ly built.
Cremohipparion - These are large to dwarf-sized hipparions; lacrimal foramen is

lacking; preorbital bar is short and lacrimal always invades the posterior portion
of the POF; when present, POF is subtriangular shaped and most usually antero-
ventrally oriented; posterior pocketing is slight to absent, medial depth is great
to slight, internal pits occur only in the most derived species, C. licenti; periph-
eral outline is strong to weakly defined, anterior rim is distinct to absent; infra-
orbital foramen is placed inferior to and encroaches upon the anteroventral
border of the POF; buccinator fossa is distinct and unpocketed except in C.
licenti; caninus fossa is present in most species, lacking only in the most primi-
tive species; malar fossa is lacking except in C. licenti; nasal notch tends to
become retracted posterior to PZ/ in this lineage; there is no persistent and
functional Pl/; maxillary cheek teeth are moderately curved to straight, have a
maximum crown height of 40-60 mm, fossette ornamentation is complex to
simple, posterior wall of postfossette is always distinct; pli caballins are variably
double or single; hypoglyphs are deeply to shallowly incised; protocones may
show some flattening lingually, but tend to become rounded, they are clearly
always isolated from the protoloph, usually have no noticeable protoconal spur,
and are lingually placed relative to the hypocone; PZ anterostyle/paraconid are
usually elongate, but become shortened in some species; mandibular incisors are
curved, not grooved, and I/3's are not atrophied; cheek tooth metaconids/meta-
stylids are rounded; ectoflexids do not separate metaconids and metastylids on
the premolars, but they do separate them on the molars; pli caballinids are
generally absent; protostylids may be present and frequent; ectostylids are
absent; linguaflexids are shallow, V-shaped to U-shaped; metapodials, when
known, are slender.
Institutional Acronyms
AL - Hadar, Ethiopian collections

AMNH- American Museum of Natural History, New York, USA
BMNH- British Museum (Natural History), London, England
BSM- Bavarian State Museum, Miinchen, West Germany
F:AM - Frick American Mammals; fossil mammal collection in the AMNH
GIU -Geologisch Institut, Utrecht, The Netherlands
GSI - Geological Survey of India, Calcutta
HLMD- Hessiches Landesmuseum, Darmstadt, West Germany
IPP- Museum of Paleontology, Sabadell, Spain
KNM - Kenya National Museums, Nairobi, Kenya
LSNK- Landessammlungen ftirNaturkunde, Karlsruhe, West Germany
MNCN - Instituto Lucas Malada, Madrid, Spain
NHMW- Naturhistorisches Museum, Wien, Austria
277
METHODOLOGY
The increased activity on hipparionine systematics in the late 1970s, and the
divergent methodologies utilized then and now, prompted the organization of an inter-
national "Hipparion Symposium." The symposium was organized by Drs. John A. Van
Couvering and Richard H. Tedford and convened by Drs. Michael 0. Woodburne and
PaulY. Sondaar at the American Museum of Natural History in the Frick Collection's
extensive equid collections. The purpose of this conference was to standardize
measurements on equid skeletons so that comparisons between investigators could be
readily made. Elements of this methodology have been utilized aJld published by a
number of authors including Eisenmann (198Z), Bernor and Hussain (1985), Bernor
(1985a), Koufos (1984, 1986, 1987a,b,c), Qiu et al. (1987), Eisenmann et al. (1988), and
Bernor et al. (1988).
Despite the plethora of literature on "Hipparion," and success in standardizing

measurements, there is still no existing consensus on species' evolutionary relation-
ships. This can be attributed to five principal factors: (1) substantial intraspecific
variability in morphological characters traditionally used; (Z) lack of attention to
stratigraphic provenance, resulting in the mixing of "population" samples often
separated by millions of years in time; (3) failure to recognize a number of evolution-
ary convergences between lineages; (4) failure to appreciate their extensive evolution-
ary diversity and long chronologie range as a group; and (5) the sheer complexity and
diversity of the samples to be studied.
These problems have been compounded by the use of divergent methodologies in

reconstructing hipparionine evolutionary series. A method popular among some
European colleagues is to collect only continuously distributed (= measured) data, and
apply numerical taxonomic methods to discover relationships (re: Eisenmann, 1981,
198Z; Koufos, 1984, 1986, 1987a,b,c). This methodology usually avoids the use of
discrete characters, develops no hypotheses of polarity, and depends mostly on distri-
bution of size and skeletal proportions to identify "evolutionary" relationships. Some-
times this methodology underestimates the number of taxa present in a sample and
cites relationship based on evolutionary grade (see Sondaar and Staesche, 1975;
Staesche and Sondaar, 1979; Sen et al., 1978). While this metholdogy is not designed
to discover phylogenetic relationships in a cladistic sense (re: Wiley, 1981), it has had
the positive effect of encouraging rigorous collection and statistical analysis of
continuous data when appropriate and justifiable. Yet, the confusion surrounding Old
World hipparionine systematics has become so great that Sondaar has stated
(1974:305): "In other words, the classification of Hipparion remains a matter of
personal taste with the various authors, because we have found no consistently distinct
morphological characters to separate species."
The last ten years have marked an effort to unravel the confusion surrounding
hipparionine evolution by developing means to identify discrete species-populations,
and characterizing them by use of both discrete and continuous (measured) charac-
ters. These attempts, as well as others, have been hindered by the fact that most
vertebrate assemblages studied contain many hipparionine species, making association
of skulls with mandibles and postcrania virtually impossible. As an example of this,
the late Miocene Greek locality of Samos has a minimum of six hipparionine species:
Hipparion dietrichi, Cremohipparion proboscideum, Cremohipparion matthewi,
"Hipparion" giganteum, Cremohipparion aff. mediterraneum, and Cremohipparion sp.
nov. (Sondaar, 1974; Woodburne and Bernor, 1980; Bernor et al., 1980; Bernor, 1985a;
Bernor et al., 1987b). These taxa have an extensive metrical overlap in size (see table
Z for a partial listing of these), which hinders accurate identification of postcrania
with specific skulls. A number of authors have elected in this circumstance to char-
acterize species on the basis of only skulls and directly associated upper cheek tooth
dentitions. There is only one circumstance that we are aware of where a single
species of hipparion is represented in a restricted stratigraphic interval by complete
articulated skeletons- the West German locality of lfowenegg. With the Howenegg
population we will be able to evaluate the range of variability of every measured and
discrete character. This will be of considerable importance in stabilizing the expected
range of variability in a single hipparionine species.
278
1) Relationship of lacrimal to the preorbital fossa:
a = lacrimal large, rectangularly b = lacrimal reduced in size, slightly

shaped, invades medial wall and invades or touches posterior border of
posterior aspect of preorbital fossa; preorbital fossa;
c = preorbitalbar (POB) long with the d = POB reduced slightly in length but
anterior edge of the lacrimal placed with the anterior edge of the lacrimal
more than 1/2 the distance from the placed still more than 1/2 the distance
anterior orbital rim to the posterior from the anterior orbital rim to the
rim of the fossa; posterior rim of the fossa;
e = POB vestigial, but lacrimal as in d; f = POB absent;
g = POB very long with anterior edge h = POB absent.

of lacrimal placed less than 1/2 the
0
distance from the anterior orbital rim
to the posterior rim of the fossa;
Z) Orbital surface of lacrimal bone:
a = with foramen; b = lacking foramen.
Fig. 1. Character states for evaluating hipparionine horse taxonomy and phylog-
eny. All characters are used for adult individuals; in addition, for cheek
tooth parameters, we refer to the state common for wear within medial 1/3
of the cheek tooth.
(continued)
279
3) Preorbital foasa morphology:
a = large, ovoid shape, antero- b = subtriangular shaped and antero-

posteriorly oriented; ventrally oriented;
c = subtriangularly shaped and d = egg-shaped and anteroposteriorly

anteroposteriorly oriented; oriented;
e = C-shaped and anteroposteriorly f =vestigial but with a C-shaped or

oriented; egg-shaped outline;
c_ -=-.)
1 = vestigial without C-shaped outline, h = elongate, anteroposteriorly oriented;
or absent;
i = morphology as in h, but with j = small, rounded structure;

faint peripheral rim;
It= posterior rim straight, with non-

oriented medial depression.
Fig. 1 (continued)
280
4) fossa posterior pocketing:
a = deeply pocketed, greater than b = pocketing reduced, moderate to

15 mm in deepest place; slight depth, less than 15 mm;
c = not pocketed but with a posterior d = absent, no rim but a remnant

rim; depression;
e =absent.
5) Fossa medial depth:
a= deep, greater than 15 mm in b = moderate depth, 10-15 mm in

deepest place; deepest place;
c =shallow depth, less than 10 mm d =absent.

in deepest place;
,....-------:::-
(~
---
,'~
=-')
6) Preorbital fossa medial wall morphology:
a = without internal pits; b = with internal pits.
7) Fossa peripheral border outline:
a = strongly delineated around b = moderately delineated around

entire periphery; periphery;
(continued)
281
c: = weakly defined around periphery; d = absent with a remnant depression;
e = absent, no remnant depression.
8) Anterior rim morphology:
a= present; b =absent.
9) Placement of infraorbital foramen:
a = placed distinctly ventral to b = inferior to, or encroaching upon

approximately 1/Z the distance between anteroventral border of the preorbital
the preorbital fossa's anteriormost fossa.
and posteriormost extent;
<©
10) Confluence of buccinator and canine fossae:
a= present; b = absent, buccinator fossa is

distinctly delimited.
11) Buccinator fossa:
a= unpocketed posteriorly; b =pocketed posteriorly.
Fig. 1 (continued)
282
lZ) Cauinus (= intermediate) fossa:
a= absent; b =present.
z--~
r::--=- ~
~ j~l'
13) Malar fossa:
a= absent;
0 ·~oo ~~
-P
14) Nasal notch position:
a = at posterior border of canine or b = approximately 1/Z the distance

slightly posterior to canine border; between canine and PZ;
c = at or near the anterior border of d = above PZ/;

PZ;
e = above P3/; f = above P4/;
g = above Ml/; h =posterior to Ml/.
~)=s-css-c;:
' I I
I I I I I I I
I I I I I I
I 1 : I I
1 I
15) Presence of P1/:
a= present, persistent and functional; b = impersistent or absent.
(continued)
283
16) Curvature of maxillary cheek teeth:
a = very curved; b = moderately curved or straight.
17) Cheek tooth crown height:
rn
a= <30 mm maximum crown height; b = 30-40 mm maximum crown height;
c = 4Q-60 mm maximum crown height; d = >6o mm maximum crown height.
\\ I
\. I
18) Maxillary cheek tooth fossette omamentaticm:
a = complex, with several deeply b = moderately complex with fewer,

amplified plications; more shortly developed plications;
t1)
c =simple complexity with few, d = generally no plis.
short plications;
19) Posterior wall of postfossette:
a = may not be distinct; b = always distinct.
Fig. 1 (continued)
284
20) P1i caballin morphology:
a= double; b =single;
c =complex; d = plis not well formed.
Zl) Hypoglyph:
a = hypocone encircled by hypoglyph; b = deeply incised, may occasionally

close, isolating hypocone;
c = moderately deeply incised; d = shallowly incised.
'-2.~
~'
ZZ) Protocone shape:
a = elongate-oval; b =lingually flattened-labially rounded;
c =oval; d =rounded;
0 0
e = triangular; f =triangular-elongate;
g = lenticular.
<::::::=::>
Z3) Isolation of protocone:
a = connected to protoloph; b = isolated from protoloph.
(continued)
285
24) Protoccmal spur:
a = elongate, strongly present; b = reduced, but usually present;
C7 a
c = very rare to absent.
0
25) Protocone/hypocone alignment:
a= anteroposteriorly aligned; b = protocone more lingually placed.
26) P2 anterostyle/paracon id:
a = elongate; b = short and rounded = - •
·~.
27) Mamh"bular in~ morphology:
a = not grooved; b =grooved.
A
28) Mandibular in~ morphology:
a= curved;. b = straight.
29) 1/3 lateral aspect:
a = elongate, not labiolingually b = labiolingually constricted.

constricted;
Fig. 1 (continued)
286
30) Premolar metaconid:
a= rounded; b = elongated; .
c = angular on distal surface.
31) Molar metaconid:
a= rounded; b = elongated;
c =angular on distal surface.
3Z) Premolar metastylid:
a= rounded; b = elongated;
c = angular on proximal surface.
~
33) Molar metastylid:
a= rounded; b = elongate;
c = angular on proximal surface.
c::)'
34) Premolar ectofiexid:
a = does not separate metaconid and b = separates metaconid and metastylid.

melastylid;
~
'JUf---/
~
a
(continued)
287
35) Molar ectofiexid:
a = does not separate metaconid and b = separates metaconid and metastylid.

metastylid;
~
'-JUf-"
~
a
36) Pli caballinid:
a= complex; b = rudimentary or single; c =absent.
37) Protostylid:
a = present, but not columnar; b = rare to absent;
\~9
c = strong, columnar.
~A
38) Ectostylids:
a= present; b =absent.
~
r ,.,r ~-1 t
3 9) Premolar linguafiexid:
a= shallow; b = V-shaped;
c =shallow U-shaped;
cb
d =deep, broad U-shape.
Fig. 1 (continued)
288
40) Molar linguafle:xids:
a= shallow; b = V-shaped;
~
c = shallow U-shaped; d =deep U-shaped.
Fig. 1 (continued)
The methodology we advocate is, whenever possible, to characterize

hipparionine taxa using both discrete and continuously distributed characters. We
attempt to characterize morphological populations as species, and using discrete
morphological characters seek to unite species into superspecific taxa using shared-
derived characters. Figure 1 contains the character states we use to define the taxa
cited here, recorded as unordered states (with no preconceived hypotheses of polar-
ity). We here evaluate competing hypotheses of hipparionine superspecific phylo-
genetic relationship citing the characters which support the various alternatives. We
then integrate this data base into a regional geochronologic framework, in order to
make some biogeographic and broader evolutionary interpretations. Here we purpose-
ly avoid using taxa that are based on limited skeletal material, i.e., cheek tooth or
postcranial material not directly associated with crania.
EVOLUTIONARY IDSTORY OF SELECTED lUPPARIONINE TAXA
We evaluate here 43 species of Old World hipparions and relevant North

American sister- and outgroup taxa based principally on the distribution of discrete
character states defined in figure 1, and recorded in table 1. We present a number of
competing hypotheses for their relationships, and summarize this information in
figures Z, 3, 6, 7, and 8.
North American Sister Taxa
North America has long been hypothesized to harbor the ancester of Old World
hipparionine horses. Recent studies by Woodburne et al. (1981), MacFadden (1984),
and Bernor et al. (1988) have particularly sought to find greater precision in the phylo-
genetic relationships between North American and Old World hipparionines. Table 1
gives the character states found in selected North American equids and Old World
hipparionines. The suite of North American equids selected for our review is believed
to be either potentially phylogenetically related to a specific clade of Old World
hipparionines, or to a suitable outgroup of all hipparionines for determining character
polarity. In an important paper by Hulbert and MacFadden (in review) many more
characters have been used than we cite here. Whereas the additional characters used
by Hulbert and MacFadden (in review) are relevant to the suite of merychippine-grade
horses that they analyze, future work is needed to determine their utility for Old
World hipparionines. We further note that many more characters are bound to emerge
with further study and comparison between Old World and New World hipparionines
and their sister-taxa, particularly as pertains to the postcranial skeleton. Our work is
merely a step in this direction.
The North American equids illustrated in our review (figure Z) include the para-
hippine horse P. leonensis (ca. 19-18 Ma; Hulbert and MacFadden, in review), M.
insignis (ca. 16=is Ma; Skinner and Taylor, 1967; Hulbert and MacFadden, in review;
289
N. A ME R I C A E U R AS I A
2 r-------~~~~~------------------~~~~----~
C. emsllel
4
C. lngenuum
6 C. pllcatlle
"H ~ Iorcel
I
8
"W proslylum \
~
9
. --
10 .
.
---=-----:::-------:c::::-.e~ "H."prlmlgen lum s.s.
" H:' tehonense
I
11
12 C. occidentale
,
-~
-
13~
~
"H:' sh lrleyl C. sphenodus

14
I
~~
15
16 ~ C.goorlsl
17
18
P. leonensis
Fig. Z. Evolutionary history of selected North American hipparionine s and the

origin of Old World hipparionine s. Reading from lower left upward, stages
of evolution of North American hipparionine s include Parahippus leonensis
(E. Hemingfordi an) as an outgroup to Merychippus insignis. M. insignis (E.
Barstovian), in turn, is considered here the sister-taxon of Cormohippa rion
goorisi (L. Barstovian), the most primitive member of the Cormohippa rion
clade, represented by the North American taxa C. goorisi, C. sphenodus
(Barstovian), C. occidentale (Clarendonia n-E. Hemphillian ), C. plicatile (L.
Clarendonia n-E. Hemphillian ), C. ingenuum (E. Clarendonia n-E. Hemphil-
lian), and C. emsliei (L. Hemphillian -Blancan). Ho: 1 (Hypothesis 1, of
Woodburne et al., 1981 and MacFadden, 1984) cites the origin of Hipparion
!•!• from a North American taxon with subsequent deployment to Eurasia,
circa 10 Ma. Ho: Z (Hypothesis Z, of Bernor, 1985a; Bernor and Hussain,
1985; Bernor et al., 1987b) cites the origin of Hipparion s •.!!_. from a primitive
"Group 1" taxon (sensu Bernor et al., 1988), and its distribution limited to
Eurasia. ---
290
Tedford et al., 1987 - which see for the approximate isotopic ages of most North
American taxa cited below; Woodburne, in preparation), C. goorisi (ca. 15 Ma;
Woodburne et al., 1981; MacFadden, 1984; Hulbert and MacFadden, in review); C.
sphenodus (ca. 14-lZ Ma; Woodburne et al., 1981; MacFadden, 1984); C. occidentale
(ca. lZ-8 Ma; Woodburne et al., 1981; MacFadden, 1984; Bernor and Hussain, 1985;
Bernor et al., 1987b, Hulbert, 1988); C. ingenuum (ca. 11-6 Ma; MacFadden, 1984;
Hulbert, 1988); C. emsliei (ca. 5-Z Ma; Hulbert, 1987, 1988); "H." shirleyi (ca. 14 Ma;
Woodburne et al., 1981; MacFadden, 1984); "H." tehonense (ca. lZ-9 Ma; MacFadden,
1984; Hulbert, 1988), and "H." forcei (ca. 11-6 Ma; MacFadden, 1984; Hulbert, 1988,
Fig. Z6). Much of this material has been recently published or is in review (Hulbert
and MacFadden, in review), and is peripheral to the focus of this paper (e.g., species of
the S\lbgenus Notiocradohipparion; C. (N.) ingenuum; C. (N.) plicatile, and C. (N.)
emsliei). Therefore, we keep our discussion of these equids to a minimum. Note that
relevant comments concerning North American "Hipparion .!•.!•" are included in the
section on Old World Hipparion s.s.
Hulbert and MacFadden (in review) have selected Parahippus leonensis as the
sister-taxon of merychippine-grade North American equids. Parahippus leonensis is a
medium sized equid with a large lacrimal that invades the medial wall and posterior
portion of the preorbital fossa (character la; table 1, figure 1). It has a lacrimal
foramen (Za). The preorbital fossa is large and ovoid in shape (3a), it is not posteriorly
pocketed, and has no posterior rim (4d), medial depth is shallow (5c), the peripheral
border is moderately well to slightly delimited (7b/c), and there is no anterior rim
(8b). The infraorbital foramen is placed distinctly ventral to the POF (9a). The
buccinator and canine fossae are confluent (lOa), with the former being unpocketed
(11a). There is no caninus fossa (lZa) nor malar fossa (13a). Nasal notch is slightly
posterior to the canine border (14a). Pl is present, persistent, and functional (15a).
The cheek teeth are very curved (16) and low crowned (17a). Although the cheek teeth
have some cement on the crown, the pre- and postfossettes have a few plications
(18c), the postfossettes' posterior walls are not consistently well formed (19a), pli
caballins are not well formed (ZOd), hypoglyphs are deeply incised (Zlb), protocones
are rounded (ZZd) but connect with the protoloph (Z3a) or in earlier wear may have an
elongate protoconal spur (Z4a), and are aligned anteroposteriorly with respect to the
hypocone (Z5a). PZ anterostyle/paraconid are rather short (Z6b).
There was an extensive middle Miocene radiation of merychippine-grade horses

in North America. Hulbert and MacFadden (in review) have evaluated some dozen of
these and verified that they are a paraphyletic group. Woodburne (in press) has noted
that of all the known merychippine horses, Merychippus insignis is the best candidate
for a sister-taxon of hipparionines. Its cranial characters are much like those of
Parahippus leonensis. The lacrimal bone has a foramen (character Za). While the
lacrimal and POF are remarkably similar in their morphology to R_. leonensis, the POF
has a posterior rim (4c), medial depth is somewhat greater (5b), and peripheral border
outline is somewhat better defined (7b). The location of the infraorbital foramen (9a),
relationship of the buccinator and canine fossae (lOa, 11a), and lack of caninus and
malar fossae (lZa and 13a) are as in P. leonensis. Likewise, the nasal notch is not
retracted far beyond the level of the-maxillary canine tooth (14a). The Pl remains
relatively large, persistent, and functional (15a). The cheek teeth have undergone a
major transformation by becoming higher crowned but remain very curved (16a) and
are still <30 mm maximum crown height (17a); pre- and postfossettes show some
simple complexity (18b/c), pli caballins have become decidedly better formed and are
usually single (ZOb), hypoglyphs may be either very deeply or deeply incised (Zla/b),
protocones are mostly rounded (ZZd), isolated from the protoloph until later wear
(Z3b), have an elongate to somewhat shortened protoconal spur (Z4a/b), protocone is
more lingually placed with respect to hypocone than seen in P. leonensis (Z5b). PZ
anterostyle/paraconid are as in P. leonensis (Z6a).
Species of the genus Cormohipparion are now under review by MOW and RLB,
and the results of that study may alter some of the interpretations reached in this
paper. Based on currently published data, Cormohipparion goori,si (MacFadden, 1984)
is about contemporaneous with Merychippus insignis. It differs in having increased
291
length of the preorbital bar and a lacrimal which may touch, but does not invade the
posteromedial wall of the POF (character 1 b). The orbital surface of the lacrimal
bone has a foramen (Za). The POF has become much better defined, it has a subtrian-
gular shape and anteroposterior orientation (3c), it is deeply pocketed posteriorly (4a),
medial depth is great (Sa), and peripheral border outline (7b) is more strongly
expressed than in M. insignis; POF has a distinct anterior rim (Sa). The infraorbital
foramen has migrated in this species to the anteroventral border of the POF (9b).
Also, the buccinator and canine fossa areas are distinct (lOb) in C. goorisi, the
buccinator fossa is unpocketed (lla), and the caninus and malar fossae are absent (lZa,
13a). The nasal notch remains unretracted in this species (14a) and Pl remains rela-
tively large (15a), but smaller than in M. insignis. The cheek teeth are similar to M.
insignis (characters 16-26), but Woodburne (personal observation) notes that they are
consistently more complex (18b/c) and higher crowned (17b). In the mandible, incisors
are not grooved (27a), and are curved (28a); the I/3's are somewhat elongate (29a);
premolar and molar metaconids and metastylids are rounded (30a, 31a, 32a, 33a);
premolar ectoflexid does not separate the metaconid/metastylid (34a), whereas in the
molars it does (35b); pli caballinids are single to rudimentary (36b); protostylids are
rare to absent (37b); ectostylids are absent (38b); premolar and molar linguaflexids are
shallow (39a, 40a).
The remaining species of North American Cormohipparion include, from oldest

to youngest, C. sphenodus, C. occidentale, C. plicatile, C. ingenuum, and C. emsliei.
The first two of these species are directly relevant to Old World hipparionine evolu-
tion and ·are discussed at length below. C. sphenodus may be a sister-taxon of Old
World "Group 1" hipparionines and Cremohipparion; the morphologic, morphometric,
and phylogenetic relatedness of C. occidentale to "Hipparion" primigenium s.s. also is
discussed, while noting that C. occidentale may contain a number of morphologies
requiring further analysis. The remaining three Cormohipparion species are relevant
only as more derived species of this genus' radiation in North America, as members of
the subgenus Notiocradohipparion Hulbert 1988, and are not considered further in this
report.
Old World •Group t• HipParionines (figure 3)
We discuss below a suite of Old World taxa in terms of some variably well-
defined groups, i.e., "Group 1," HipParion ~·~·• the "Sivalhippus Complex," and
Cremohipparion. Some of these groups are currently defensible as "natural," or mono-
phyletic, others are clearly paraphyletic, while yet others require much further study
and analysis to stabilize their taxonomy, and to understand their phylogenetic rela-
tionships.
The most primitive group, and most likely one that would appear to have had its
origins from a North American Cormohipparion taxon is the "Group 1" assemblage
presented here in figure 3. Woodburne and Bernor (1980:1328) initially defined this
group as being: " ••• composed of hipparionine horses with a large, well defined, pre-
orbital fossa separated from the orbit by a relatively great distance (= preorbital
bar)." They made a number of observations on the preorbital fossa, cheek tooth, and
postcranial morphologies, when information was available, and commented on its
likely derivation from North American Cormohipparion, following Skinner and
MacFadden (1977). Bernor et al. (1980) applied what was then understood of "Group 1"
evolutionary relationships, along with other hipparion lineages, to an early Old World
biochronologic ranking of hipparionines. Subsequently, Woodburne et al. (1981) evalu-
ated the phylogenetic relationships between Old World "Group 1" horses, the chronol-
ogy of the North American and Old World "Hipparion Datum," and specifically the
viability of earlier proposals that North American Cormohipparion was related to
"Hipparion" primigenium. Further systematic work by Bernor and Hussain (1985) and
Bernor (1985a) have refined the systematics of this group. Bernor et al. (1988) found
it to be paraphyletic, while Woodburne (in press) has further considered the relation-
ships of .the group to North American hipparionines. We draw upon this literature
here.
292
M W EUROPE E. MEDITERRAN EAN, S.W. ASIA CHINA N. AFRICA
6
large sp. small sp.
,
' I
', I
\ I
\ I
II
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
Fig. 3. Evolutionary history of primitive Old World "Group 1" (sensu Bernor et
al., 1988) hipparionines; "Hipparion" primigenium s.s. is potentially the
ancestor of a number of distinct Old World clades; in western Europe,
the "H." catalaunicum-"H. mediterraneum" clade and the "H."
melendezi clade; in the western U.S.S.R., eastern Mediterranean, and
southwest Asia, the H. ~-H. prostylum clade (Hipparion s._!.; Ho: 2
presented here in figures 2. and 6) and the "H." giganteum clade; in
North Asia, the "H. weihoense-"H." platyodus-"H." coelophyes-"H."
hippidiodus transformation series (Bernor et al., in press) and the "H."
dermatorhium clade.
Taxa which we include in "Group 1" are (see definitions given above) "Hipparion"
primigenium .!•.!• (ca. 11-?8 Ma; Tobien, 1986; Bernor et al., 1987b, 1988, in press);
"Hipparion" catalaunicum (ca. 11 Ma; Woodburne and Bernor, 1980; Bernor and
Hussain, 1985; Bernor et al., 1987b, 1988); "H. mediterraneum" (Alberdi, 1974); "H."
melendezi (ca. 9.5 Ma; Alberdi, 1974; Woodburne and Bernor, 1980; Bernor and
Hussain, 1985; Bernor et al., l987b, 1988); "H." gettyi (Woodburne and Bernor, 1980;
Bernor, 1985a; Bernor et al., 1987b, 1988); "H." giganteum (ca. 8 Ma; Gromova, 1952.;
Gabunja, 1959; Woodburne and Bernor, 1980; Bernor et al., 1987b, 1988); "Hipparion"
africanum (ca. 10.5 Ma; Arambourg, 1959; Woodburne and Bernor, 1980; Eisenmann,
1979; Bernor et al., 1987a,b, 1988); "Hipparion" weihoense (ca. 11-10 Ma; Liu et al.,
1978; Bernor et al., 1988, in press); "H." dermatorhinum (ca. 1Q-7.5 Ma; Bernor et al.,
1987b, 1988, in press; Qiu et al., 1987); "H." coelophyes (ca. 8 Ma; Bernor et al., 1987b,
1988, in press; Qiu et al., 1987); "H." hippidiodus (ca. 8-6 Ma; Bernor et al., 1987b, in
press; Qiu et al., 1987).
"Hipparion" primigenium .!·.!• (Bernor et al., 1988) is ostensibly the most primi-
tive Old World member of this group morphologically, and chronologically the oldest
Eurasian hipparion. As such, it occupies an important place in Old World hipparionine
phylogenetic& in that, beyond its morphological and chronological attributes, it may
represent the sister taxon for all Old World hipparionines (Bernor and Hussain, 1985;
Bernor et al., 1988). "Hipparion" primigenium .!·.!· is known from Central Europe,
having first appeared circa 11.5-11 Ma, and its morpholgy is best recorded in horizons
of Pannonian age in the Vienna Basin (NHWM collections, Bernor et al., 1988) and at
293
the German locality of Kowenegg (LSNK and HLMD collections, Tobien, 1986; Bernor
et al., in press). It is characterized as being a large hipparionine, with a long
preorbital bar and the lacrimal clearly placed more than one-half the distance from
the anterior orbital rim to the posterior rim of the fossa (table 1, character lc). As
with all Old World hipparionines, as far as we know, the orbital surface of the lacrimal
bone lacks a foramen (2.b). The preorbital fossa is subtriangular shaped and antero-
ventrally oriented (3b); it is deeply pocketed (4a), being greater than 15 mm in its
deepest place; its medial depth is great (Sa), being more than 15 mm in its deepest
place; its medial wall lacks internal pits (6a); it has a strongly delineated peripl!.eral
border outline (7 a); it has a distinctly defined anterior rim (Sa); the infraorbital
foramen is placed inferior to, or encroaches upon the anteroventral border of the
preorbital fossa (9b). The buccinator and canine fossae are distinct (lOb); the
buccinator fossa is not posteriorly pocketed (lla); there is no caninus fossa depression
(12.a) and there is no malar fossa (13a). The nasal notch is retracted to about the
anterior border of PZ/ (14c). When present, Pl/ is very reduced in size and clearly
nonfunctional (15b). The maxillary cheek teeth are moderately curved to straight
(16b); unworn cheek teeth have a maximum crown height between 40 and 60 mm (17c);
enamel plications of the pre- and postfossettes are complex (18a), maintaining several
deeply amplified plis until well beyond the middle stage-of-ear; the posterior wall of
the postfossette is always distinct (19b); pli caballins are mostly double (ZOa); hypo-
glyph is deeply incised well beyond the middle stage-of-ear (Zlb); protocone is
commonly lingually flattened and labially rounded (2.Zb; some approaching ZZc, nearly
oval, depending upon wear-stage and population sampled) and is clearly isolated from
the protoloph until very late stage-of-ear (2.3b; if ever); the protocone spur is very
rare to absent (Z4c); the protocone is more lingually placed than the hypocone (ZSb);
P2. anterostyle/paraconid is elongate (2.6a). The mandibular incisors are not grooved
(Z7a), they are curved (ZSa), and I/3 is elongate but not transversely constricted (Z9a);
premolars and molars have rounded metaconids and metastylids (30a, 3la, 3Za, 33a);
the premolar ectoflexids do not separate the metaconid and metastylid (34a), although
they do so in the molars (35b); lower cheek teeth frequently have complex to single pli
caballinids (36a/b); protostylids are commonly present, but never strong and cohlmnar
(37a); ectostylids are rare to absent (38b); linguaflexids are usually shallow on the
premolars (39a), while being more V-shaped on the molars (40b).
The metapodials of ".!!:" primigenium have commonly been characterized as

being robust (Forsten, 1968; Woodburne and Bernor, 1980). Bivariate plots comparing
the Howenegg sample's MC m and MT m maximum length (variable 1) and distal
articular width (variable 11) with those of the Xmas Quarry sample of Cormohipparion
occidentale (sensu MacFadden, 1984) are given in figure 4 ... These comparisons show,
by use of 95% and 99% confidence interval ellipses of the Howenegg sample, that the
range of variation in the Xmas Quarry sample far exceeds those in the Iiowenegg
sample. In those figures where ellipses of the Howenegg sample are superimposed
over the Xmas Quarry plots, at least three populations can be discerned. MacFadden
(1984) observed morphological variation in Cormohipparion occidentale skulls from
Hans Jolmson Quarry, and hypothesized that this variability was due to sexual
dimorphism. If the bivariate plots we figure here are an accurate gauge of "Cormo-
hipparion occidentale"'s variability, then a number of subtly different, but in many
ways morphologically similar, species may now be included within the taxon "C.
occidentale". This point of view was also expressed by Eisenmann et al. (1987) based
on a quantitative analysis of skull proportions.
The extent of morphological similarity between "Hipparion" primigenium s.s. and

"Cormohipparion occidentale" (~ lato) is remarkable, as can be seen in comparing
the character states of the species given in table 1 and as rendered here in figure 5
(also compare descriptions of MacFadden, 1984 and Bernor et al., 1988). If, however,
material now identified as Cormohipparion occidentale is comprised of more than one
species, it remains to be seen which of these species, if any, is the most likely
ancestor for "Hipparion' primigenium !.•!.• At least, a detailed morphologic and
morphometric study of this problem is needed to revise the alpha-taxonomy of "C.
occidentale" and determine which of several morphs may be referred to this species,
and which of these morphs is morphologically most similar to "Hipparion" primigenium
294
Hoawenagg HiPPer ion XMAS ClJarry COriiOhipperion
MCII! MC III
270 270
I I I
A B
240 240
...
~99%EIIipse
.·
210 ~95%EIIipse 210
] ]
• lBO • lBO
150 150
120 I 120 I I I
15 23 31 39 47 55 15 23 31 39 47 55
DAM (-.! DAM (ooJ
... 0.
Hoewenegg Hipparion XMAS QJarry COriiOhipperion
MT Ill MT Ill
270 270 I
I I I I I
c D
~99%EWpse
99%EIIipse
-.". .
240 240 • •• • • 95% Ellipse
95% Ellips•
210 210
:
] j
• lBO • lBO
150 150 .,
I I ( I
120 120
15 23 31 39 47 55 15 23 31 39 47 55
DAM (ooJ DAM (ooJ
Fig. 4. Bivariate plots comparing metapodial dimensions of "Hipparion"

r:rimigenium (Howenegg) and North American Cormohipparion occidentale
Xmas Quarry). A. Bivariate plot of Howenegg hipparion MC m maximum
length (variable 1) X distal articular width (variable 11) with 95% and 99%
confidence interval ellipses drawn in relationship to individual observed
measurements. B. Bivariate plot of Xmas Quarry "Cormohipparion" MC m
measurements 1 X 11 with 95% and 99% Howenegg confidence interval
ellipses superimposed. This plot shows that a minimum of three populations
are present within the Xmas Quarry sample. C. Bivariate plot of Howenegg
hipparion MT m measurements 1 X 11 with 95% and 99% confidence inter-
val ellipses drawn in relationship to observed measurements. D. Bivariate
plot of Xmas Quarry "Cormohipparion" MT m measurements 1 X 11 with
95% and 99% Howenegg confidence interval ellipses superimposed. Note
that once again a minimum of three Xmas Quarry populations is suggested.
However, the largest cluster would appear to have a variance greater than
the Howenegg sample and may represent two separate, but similarly sized
populations.
295
Fig. 5. Morphological comparison of North American Cormohipparion
occidentale and central European "Hipparion" primigenium !.·~
A. Cormohipparion occidentale F:AM 71800, from Xmas Quarry,
Ash Hollow Formation, late Clarendonian age (ca. 9 Ma), of
Cherry County, north~entral Nebraska. Left lateral view of
skull. B. F:AM 71800, left occlusal view of PZ/-M1/. C. F:AM
71800, left occlusal view of P/Z-M/3. D. "Hipparion" primigenium
'!.·!.· LSNK lfo A, from lfowenegg, early Vallesian age (ca.
10.8 Ma), Hegau, West Germany. Left lateral view of skull.
E. LSNK lf'o A, left occlusal view of PZ/-M3/. F. LSNK Ho A, left
occlusal view of P/Z-M/3. G. "Hipparion" primigenium !.·!.-
NHMW 17, from Gaiselberg, earliest Vallesian (ca. 11.5-11 Ma),
Vienna Basin, Austria. Right M1/ (reversed). Note here the
similarities of skull, maxillary cheek tooth, and mandibular cheek
tooth character states after descriptions in text and table 1. Also
note (G) the Cormohipparion-like protocone on some specimens of
the chronologically oldest European hipparion from Gaiselberg,
Vienna Basin (see Bernor et al., 1988 for further discussion of the
Vienna Basin population of "Hipparion" primigenium !.·!1·
Figures SA, B, and C were redrafted from MacFadden (1984) and

adjusted here to scale equivalently to the lfowenegg horse. Figure
SG is twice the size of equivalent M1/ rendered in SB and SE.
!.·!.·The illustration of this particular problem further shows the value of the
H~wenegg quarry sample and the need to utilize parametric statistics for alpha-level
taxonomic work.
Of the "Group 1" hipparions considered here, "H." w eihoense is the most similar
to "H." primigenium in its morphology (Bemor et al., 1988), but may differ in a more
anteroposterior orientation of its POF (character 3c), a more elongate snout and,
perhaps, more elongate protocones (ZZa) (Woodburne, in press). In light of these
potential differences and the fact that postcranial elements still have not been
reported for the Chinese form, we retain it as a species distinct from "H."
primigenium s.!_.
"Hipparion" catalaunicum, from Hostalets Superior, near Barcelona, Spain, is

likewise a large horse, virtually identical in size to "H." primigenium s.s. in the shape
of its preorbital fossa, which is very elongate, but yet oriented anteroposteriorly
(character 3h). Neither an associated mandible nor postcranial remains have been
reported for this taxon, making comparisons to "H." primigenium s.s. impossible. The
Bou Hanifia, Algeria, horse "Hipparion" africanum (Arambourg, 1959; Eisenmann,
1979; Woodburne and Bernor, 1980; Bernor et al., 1987a) shows all the same character
296
states of the skull that "H." catalaunicum does (table 1) except for somewhat more
retracted nasals (14d); protocones show some lingual flattening but are more elongate-
oval shaped (ZZa/b); pli caballinids are variably complex or single (36a/b); protostylids
are not prominent, but are present (37a); cheek tooth linguaflexids are V- to shallow
U-shaped (39b/c, 40b/c). "H." africanum also has associated mandibular and post-
cranial remains. Whereas mandibular morphology is virtually the same in this species
as in "Hipparion" primigenium, "H." africanum has decidedly shorter and more gracile
metapodials (see table Z here and Bernor et al., 1987a).
Alberdi (1974) reported, but did not figure, an important skull of "Hipparion
mediterraneum" from the Turolian locality of Piera, near Barcelona, Spain. This
specimen is not attributable to "H." mediterraneum as recognized by Bernor (1985a)
(and here Cremohipparion mediterraneum), but rather shows a remarkable resem-
blance to the stratigraphically older taxon, "H." catalaunicum. The facial morphology
is very reminiscent of "H." catalaunicum in its elongate and anteroposteriorly directed
POF morphology (3i). However, the POF is relatively derived in its absence of poste-
rior pocketing (4d), shallow medial depth (5c), weakly defined peripheral border (7c)
and lack of an anterior rim (8b). The cheek teeth also appear to have undergone some
simplication of fossette complexity (18b) and had somewhat less deeply incised hypo-
glyphs (Z1b), and protocones had become more rounded (ZZd; Alberdi, 1974). It would
appear that the Piera hipparionine may have been an endemically evolved species,
derived from "Hipparion" catalaunicum.
"Hipparion" ~ originates from the lower biostratigraphic interval at

Maragheh (Bernor, 1985a, 1986). It is decidedly smaller than "Hipparion" primigenium
.!·.!•• falling well below its observed range for PZ/-M3/ (table Z). Likewise, it is derived
compared to "H." primigenium in that the anterior rim of the preorbital fossa has been
rotated dorsally, giving the fossa a subtriangular, anteroposteriorly oriented shape
(character 3c); preorbital fossa medial depth is relatively reduced (5b); peripheral
border outline is only moderately delineated (7b) and the anterior rim is lacking (8b).
All of these features indicate an initial reduction of the preorbital fossa. The cheek
teeth also show some derivations compared to "Hipparion" primigenium s.s. including
less complex plications of the fossettes (18b); the hypoglyph is somewhat less deeply
incised (Z1c); protocone has a more oval morphology (ZZc). The rectangular shape of
the lacrimal in this species (figure 3) differs from virtually all other hipparionines.
Interestingly, the lacrimal shape in "H."~ is similar to that in "H."
mediterraneum" (figure 3). This warrants further study. Bernor (1985a) also reported
a rather elongate and slender metapodial for "H." ~ but recent reevaluation of
the NHMW collection (by RLB) has revealed that a second horse, "Hipparion" aff.
moldavicum, is known from the lower stratigraphic interval as well, making any
postcranial and isolated dental referrals to "H." gettyi uncertain. However, the
morphology of the type of "Hipparion" gettyi is distinctly divergent from "Hipparion"
primigenium .!•.!• and can be interpreted to trend in the direction of Old World
Hipparion .!•.!• species. This is the reason why Bernor and Hussain (1985) and Bernor
(1985a) advanced the hypothesis that the Hipparion s.s. lineage had an Old World origin
and geographic distribution. This is discussed further below.
"Hipparion" melendezi from late Vallesian horizons of Castilla la Vieja (Alberdi,

1974: Plate 1) is the same size as "Hipparion" gettyi. It is derived relative to "H."
primigenium !•!• in its smaller size, the very reduced length of its subtriangular
shaped, anteroventrally oriented preorbital fossa (3j); reduced fossa medial depth (5b),
peripheral border outline (7b), and absent anterior rim (8b). In the cheek teeth only
hypoglyphs would appear somewhat reduced in their incision (Z1b/c). This lineage,
currently recognized as monospecific, would represent a clade separate from the "H."
catalaunicum-"H. mediterraneum" lineage. -
"Hipparion" giganteum has virtually the same size PZ/-M3/ dimensions as

"Hipparion" primigenium .!·.!·• but is remarkable for its long and robust snout. It is
medial Turolian (= Meotian), and has been recorded from Samos, as well as a number
of western U.S.S.R. localities (Gromova, 195Z; Gabunja, 1959; "Hipparion" sp. of
Woodburne and Bernor, 1980; Bernor, 1985a). "H." giganteum appears to be derived
297
directly from "H." primigenium !•!•' having few morphological character state trans-
formations including snout lengthening, consistently shorter preorbital fossa
(Woodburne and Bernor, 1980); reduced preorbital fossa pocketing (character 4b),
moderate medial depth of the fossa (5b), lack of a POF anterior rim (8b), and some-
what reduced hypoglyph incision (Z1c). Unfortunately, only some western U.S.S.R.
material has associated mandibular and postcranial remains, and Gabunja's measure-
ments are not currently available to us (his tables are not translated from Russian).
Gabunja (1950: 17-18) has included in his diagnosis of this taxon that the metaconid and
metastylid are of the "hipparion type" (i.e., rounded; characters 30a, 31a, 3Za, 33a)
with massive metapodials, and weakly developed lateral digits.
"Hipparion" dermatorhinum is a large, highly derived hipparion reported from

China (Sefve, 1927; Bernor et al., 1987b, in press; Qiu et al., 1987) as well as Central
Asia (Zhegallo, 1978). While having virtually the same length PZ/-M3/ as "H."
primigenium !•!•' this taxon is derived in a number of cranial features. "H."
dermatorhinum has an elongate slender snout with the anterior dentition having a
mediolaterally narrow arcade-shaped semicircular contour. This species has a slightly
reduced preorbital bar, and the lacrimal bone does not contact the posterior extent of
the POF (character 1d). The fossa's posterior pocketing is sharply reduced (4b) and its
medial depth is shallow (5b). The nasal notch is sharply retracted to a position above
P4/ (14f). Protocone morphology of "Hipparion" dermatorhinum is unique for Old
World hipparionines in its elongate, lenticular shape (ZZg). In other cheek tooth char-
acters, "H." dermatorhinum is conservative and similar to "H." primigenium !·!·
Mandibular cheek tooth characters are, for the most part, as in "H." primigenium s.!.,
with the remarkable exception that the metaconids and metastylids tend to be quite
elongated (30b, 31b, 3Zb, 33b). Molar linguaflexids are deeper and U-shaped. Directly
associated postcrania have not been reported for "H." dermatorhinum.
The last two taxa referred to "Group 1" are the Chinese taxa "Hipparion"
coelophyes and "Hipparion" hippidiodus. Bernor et al. (1987b) and Bernor et al. (in
press) have argued that these species represent the final two steps of a Chinese
morphological transformation series, although they are distinctly not chronospecies.
They are similar sized, and the smallest "Group 1" taxa. "H." coelophyes is derived
compared to "H." primigenium !•!• in its somewhat more egg-shaped, and antero-
posteriorly oriented preorbital fossa (character 3d). Also, the preorbital fossa is not
pocketed, but retains a strongly developed posterior rim (4c); the fossa is reduced
somewhat in its medial depth (5b/c); the peripheral border outline is only moderately
well delineated (7b/c) and the anterior rim is absent (8b). Despite these facial diver-
gences, the cheek teeth are completely primitive in their morphology. "Hipparion"
hippidiodus is further derived in its near to complete loss of the preorbital fossa (1e,
3g, 4d, 5c, 7d). Also, in comparison to "H." coelophyes, the cheek teeth are simpler in
their fossette ornamentation (18b/c); pli caballins are single (ZOb), hypoglyphs are
more shallowly incised (Z1c/d), and protocone has become round (ZZc/d) by losing its
lingual flatness. In all these regards, "Hipparion" hippidiodus parallels Hipparion s•!• in
its transformation of facial and cheek tooth characters (see the next section for
further discussion of this problem).
Old World Hipparion s.s.
The concept of Hipparion !·!· is based on the species Hipparion prostylum

Christo! (183Z) from the medial Turolian locality of Mt. Luberon (= Cucuron, after the
adjacent town, or Mt. Leberon, an alternate name used for the mountainside where the
deposits are located), Province of Vaucluse, France (MacFadden, 1980; Woodburne and
Bernor, 1980). Christo! noted that this horse was characterized (in part) by three toes
on each foot and an isolated protocone. In 1849, Gervais reiterated these two charac-
teristics and further mentioned the occurrence of additional stylids on lower deciduous
molars. Gervais (1849) designated the three Mt. Luberon species; H. prostylum, H.
diplostylum, and H. mesostylum. Later, Gervais (1859) united all of these species into
H. prostylum, recognizing the ontogenetic variability of these characters. Gaudry
U873), Osborn (1918), and in later years Sondaar (1974), Skinner and MacFadden
(1977), Woodburne and Bernor (1980), MacFadden (1980, 1984), and Bernor (1985a) have
298
all agreed in referring at least the majority of the Mt. Luberon material to H.
prostylum. Sondaar (1974) stated that the specimen figured by Gervais (1859; pl. 19,
fig. Z; maxilla fragment with P3/-MZ/) is the holotype of H. prostylum. Bernor (1985a)
noted that a type was never assigned, -and rectified the problem by assigning the most
complete specimen, BMNH M33603, to H. prostylum. This action also maintained the
original concept of H. prostylum as much as possible.
Sondaar (1974) stated that some postcranial elements in the Mt. Luberon
hipparion sample indicate a larger rare species. Following an_ unpublished manuscript
by Woodbume, Bernor (1985a) noted the existence of two cranial morphologies, one a
rare form with a better developed facial fossa (BMNH MZ6617; Figs. Z, 3, and 6) which
he referred to H. aff. prostylum, and the other, more common form which he referred
to H. prostylum. It would be tempting to associate the rare larger postcrania to H.
aff. prostylum, and the remaining majority of postcrania to H. prostylum s.!_., but this
solution, if viable, depends on further analytical work underway by Samir in Paris.
Nevertheless, Woodbume (in Bernor et al., 1980) maintains that the rare second
species likely is representedby BMNH MZ6617 which, with a large and anteroventrally
oriented POF and large preorbital bar, is a member of "Hipparion" Group 1.
These definitions of the genotype species Hipparion !·.!· are critical for evalu-
ating the varying hypotheses about its phylogenetic and biogeographic relationships.
MacFadden (1980, 1984), Woodburne (in press), and Hulbert and MacFadden (in press)
support the hypothesis that Hipparion .!•.!• has its origins in North America, the most
primitive species being H. shirleyi (ca. 14 Ma), from late Barstovian age horizons of
Polk County, Texas Gulf Coastal Plain (MacFadden, 1984). Other North American
taxa include H. tehonense (lZ-9 Ma) and H. forcei (11-6 Ma). Bernor and Hussain
(1985), Bernor(l985a), and Bernor et al. (1987b) have argued for an Old World origin of
Hipparion .!•.!•• citing apparent convergences in facial characters between North
American forms and Old World Hipparion s.s. There is little disagreement about which
Old World species are referable to Hipparion .!•!•• only their extra-Eurasian phylo-
genetic relationships. We return to this issue below, following the description of Old
World members of Hipparion s._!.
Members of Old World Hipparion s._!. recognized here (figure 6; same as Group 3
hipparions of Woodbume and Bernor, 1980 and Bernor, 1985a) include H. prostylum (ca.
8.5 Ma; Woodbume and Bernor, 1980; Bernor et al., 1980, 1987b; Bernor, 1985a); H.
dietrichi (ca. 8-7 Ma; Woodburne and Bernor, 1980; Bernor et al., 1980, 1987b; Bernor,
1985a); H. campbelli (ca. 7 Ma; Bernor, 1985a); H. antelopinum (late Miocene;
Woodburne and Bernor, 1980; MacFadden and Woodburne, 198Z; Bernor and Hussain,
1985; Bernor et al., 1987b).
Hipparion prostylum is a medium sized hipparion. It has a preorbital bar that is

slightly reduced in length but with the anterior edge of the lacrimal placed still more
than one-half the distance from the anterior orbital rim to the posterior rim of the
fossa (character ld). H. prostylum lacks an orbital foramen (Zb). It has an egg-shaped
and anteroposteriorly oriented preorbital fossa (3d); POF posterior pocketing is
reduced and variably of moderate depth or no measurable depth, having a well deline-
ated posterior rim (4b/c), medial wall depth is moderate (Sb), lacks internal pits (6a),
is moderately to weakly defined around the periphery (7b/c), and the anterior r m is
generally absent (8b). The infraorbital foramen usually encroaches upon the antero-
ventral border of the preorbital fossa (9b); the buccinator fossa is separate from the
area of the canine fossa (lOb) and unpocketed posteriorly (lla); there is neither a
caninus fossa (lZa) nor a malar fossa (13a). The position of the nasal notch is at or
near the anterior border of PZ/ (14c). Pl/ is usually not present, but it is nonpersist-
ent and nonfunctional when present (15b). Maxillary cheek teeth are generally
straight (16b), unworn cheek teeth have a maximum crown height of about 50 mm
(17c); pre- and postfossettes are moderately complex, often having many plis, but of
markedly shortened amplitude (18b); posterior wall of postfossette is distinct (19b); pli
caballines are variably double or single (ZOa/b); hypoglyphs are deeply to moderately
deeply incised (Z1b/c); protocones are oval-shaped structures (ZZc); protocone is
isolated from protoloph until very late stage-of-wear (Z3b); protoconal spurs are very
rare to absent (Z4c); protocones are more lingually placed than the hypocone (ZSb). PZ
299
M EU RO PE N. ME OITERRANEAN S. A 5 I A
. ~~
~ I ~ ~-.(. H. campbelll
~
---
8 H . dietrichi __..,....-
~ "'""'~···
9
~ / pm•tylum
10
r
,, ~
12
"H:' primlgenlum
\
'"H~ shirley!
Fig. 6. Evolutionary history of Old World Hipparion s.~. From the lower right-
hand corner, Ho: 1 (Hypothesis 1 of Woodburne et al., 1981) Hipparion
.!!.•.!• from a North American ancestor close to Hipparion shirleyi; Ho: 2
(Hypothesis Z of Bernor, 1985a; Bernor and Hussain, 1985, and Bernor
et al., 1987b), the origin of Old World Hipparion s.s. from a population
of "H." primigenium .!•.!• Both hypotheses support the subsequent
evolution of more derived eastern Mediterranean and southwest Asian
species H. dietrichi (M. Turolian) and H. campbelli (M. Turolian), and
south Asian species H. antelopinum (L. Miocene).
anterostyle/paraconid are elongate (26a). The postcranium includes elongate meta-

podials. The Mt. Luberon sample of mandibles include the following dental features:
incisors are not grooved (27a), are curved (28a), and are not transversely constricted
(29a); cheek tooth metaconids and metastylids are usually rounded (30a, 31a, 32a; 33a);
premolar ectoflexids do not separate metaconid and metastylid (34a), whereas they do
in the molars (35b); pli caballinids are rudimentary or single (36b); protostylids may or
may not be seen occlusally, but a mandible in Paris (MNHN 101) has the cement
eroded labially, and shows that while the protostylids are not always expressed occlu-
sally they were strongly developed beneath (37b); ectostylids are absent (38b); pre-
molar and molar linguaflexids are variably V- to shallow U- shaped (39b/c, 40b/c).
Hipparion dietrichi and Hipparion campbelli both differ from Hipparion

prostylum in that their facial fossae are reduced to a C-shaped or egg-shaped outline
(character 3f); H. campbelli retains a very small sharp segment of posterior r m,
whereas H. dietrichi has a thickened ledge (both 4c/d); both species have medial depth
reduced to a shallow state (Sc) with a remnant depression. Whereas H. dietrichi's
nasal incision is as in H. prostylum (14c), H. campbelli's is retracted slightly more to a
position above the mesostyle of PZ/ (14d). Hipparion campbelli has both associated
lower jaws and postcranial remains. In the mandible, incisors are not grooved (Z7a),
curved (Z8a), and 1/3 is elongate, not transversely constricted (30a); premolar and
molar metaconids and metastylids are rounded (30a, 31a, 32a, 33a); premolar ecto-
flexids do not separate metaconid and metastylid (34a), whereas they do in the molars
(35b); pli caballinids and ectostylids are absent (36c; 38b); protostylids are rarely
300
present (37b); linguafiexids are shallow on premolars (39a), and deeper on the molars
(40c). Hipparion campbelli and Hipparion dietrichi differ significantly from one
another; the former has a narrow snout with anterior dentition arranged in a narrow
arcade-shaped semicircle, the latter having a broa<l incisor region with incisors all
anteriorly placed. Solounias et al. (1988) have made studies comparing Miocene
giraffid material with extant ruminants and have found that the morphological pattern
exemplified by H. campbelli is associated with selective, more browsing regimes,
whereas the pattern exemplified by H. dietrichi is typical of animals adapted to
grazing regimes. This potential morphological and dietary relationship serves to
further distinguish H. campbelli from H. dietrichi.
Hipparion antelopinum is not very well represented by fossil material. The type
assemblage, preserved in the BMNH, is small and fragmentary (e.g., MacFadden and
Woodburne, 198Z, present additional discussion and illustrations of this material).
There is a skull, BSM 689 located in Munich, which preserves more features, but its
referral to this taxon is uncertain (see Bernor and Hussain, 1985, for a discussion of
this specific problem). However, if one were to include this specimen, there is little
to deny it being an advanced member of Old World Hipparion s.s. It is most similar to
H. dietrichi and H. campbelli in its reduced preorbital fossa morphology (characters
l?d, Zb, 3?c, 4?c, 5c, 6a, 7?c, 8b). In the cheek teeth the type series of H.
antelopinum has a moderately complex enamel plication pattern (18b), single pli
caballins (ZOb; whereas the Munich specimen has persistent double pli caballins, ZOa),
moderately deeply incised hypoglyphs (Z1c) and oval protocone (ZZc). The type collec-
tion preserves some metapodials which are rather elongate and slender. Associated
lower cheek teeth have the same suite of morphological characters as H. campbelli.
MacFadden (1980, 1984) and Woodburne (in press) have advanced the hypothesis
that Hipparion s.s. originated in North America. MacFadden (1984) has referred three
North American species to Hipparion !•!• including Hipparion shirleyi (14 Ma),
Hipparion tehonense (1Z-9 Ma), and Hipparion forcei (11-6 Ma). There are a constella-
tion of characters within the diagnoses for these hipparionines which are consistent
with an Hipparion !•!• referral including a preorbital fossa which is usually anterior to
the lacrimal, placed high on the face, which is only moderately deeply pocketed poste-
riorly, only moderately deep medially, has no anterior rim and has only a moderately
well delineated peripheral rim; the nasal notch is incised to a level of PZ/; protocone&
are oval shaped; the lacrimal is subtriangular shaped; metapodials are rather elongate
and slender.
In comparing these North American taxa with Old World Hipparion s.s., we list a
number of characters which do not support this phylogenetic hypothesis, particularly
in Hipparion shirleyi (which is the most primitive North American taxon; holotype
F:AM 79950). These characters include: lacrimal contacts the preorbital fossa (char-
acter 1b); the infraobital foramen is placed above P4/, along the ventral border of the
preorbital fossa, instead of at its anteroventral rim (9a); P1/ is relatively large, and
apparently functional (15a); cheek teeth are quite low crowned, being only Z5-30 mm
in unworn mesostyle height (17a); cheek teeth have a very simple complexity of the
fossette borders (18c); posterior wall of the postfossette is not always distinct and
well formed (19a); pli caballins are lacking altogether on PZ/ (ZOd), while being single
on all other molars (ZOb); cheek tooth hypoglyphs are very deeply incised, sometimes
encircling the hypocone altogether (Z1a); protocone has a well developed anterior spur
(Z4b); anterostyles/paraconids are short (Z6b); ectoflexids are deep on the premolars,
separating metaconid and metastylid (34b); size is significantly smaller than any Old
World Hipparion s.s.
MacFadden (1984) includes two more derived species in North American

Hipparion !•!·= Hipparion tehonense and Hipparion forcei. Hipparion tehonense is
more similar to Old World Hipparion !•!• than H. shirleyi in overall cranial size,
anterior orientation, and decreased depth of the POF, yet differs in retaining a
functionally persistent P1/ (character 15a); it has simpler plication& of the cheek tooth
fossettes (18c); PZ/ anterostyle is short (Z6b). Hipparion forcei is most like Old World
Hipparion !•!•' differing in a preorbital fossa morphology that may have an obliquely
301
disposed, very straight posterior rim and non-oriented medial depression (character 3k)
in the referred, crushed cranium from California (MacFadden, 1984). MacFadden
(1984) also refers specimens to this species from the late Clarendonian of Nebraska
that show a more normally-appearing POF (3c); enamel pliations are moderately
complex (18b/c) in both samples, but not quite as complex as in H. prostylum or H.
dietrichi which commonly are more strongly plicated on all fossette margins, but
especially on the anterior border of the prefossette and posterior border of the post-
fossette.
The hypothesis favored by MacFadden (1984) and Woodbume (in press) considers
that virtually all of the incongruent characters found in H. shirleyi are plesiomorphies,
and that the transformations required to develop morphologies displayed by either
North American or Old World species would be comparable in either case. In fact, it
appears that these kinds of transformations (e.g., increased length of the preorbital
bar, with concomitant restriction of the lacrimal from the rear of the POF; increase
in general size, in hypsodonty, in dental complexity, have occurred independently, and
thus convergently, in other equid lineages that began from a "merychippine grade"
ancestor. Neohipparion, ~MacFadden (1984) is one such example. Thus, while a
detailed transformation series to demonstrate the progressive acquisition of the
above-cited changes is not now known, the proposal that Hipparion shirleyi stands at
the base of Hipparion s.s. is plausible.
On the other hand, Bemor (1985a, 1986) and Bemor and Hussain (1985) propose
that Hipparion s.s. had an endemic origin in the Old World and that North American
species now referred (MacFadden, 1984) to that genus were derived convergently. In
support of an Old World origin of Hipparion .!•.!•' one finds biostratigraphic and
geochronologic congruence. Bemor's (1985a) proposal of potential phylogenetic rela-
tionships between "Hipparion" gettyi and Hipparion prostylum is represented in a
single section ·of rocks, calibrated at Maragheh, Iran (Bemor, 1986). Furthermore,
taxa resembling both "Hipparion" gettyi and Hipparion prostylum are found at the
localities of Mt. Luberon as well as Pikermi (Bemor, 1985a). Bemor (1985a) referred
BMNH ZZ617 from Mt. Luberon to Hipparion aff. prostylum, whereas Woodbume (in
press) believes that it pertains to a Group 1 "Hipparion." The reader should keep these
complexities in mind when surveying figures 3 and 6. The specimen labeled there as
Hipparion prostylum is BMNH ZZ617. In any case, Hipparion prostylum is known from
a number of localities throughout southwestern Europe, the eastern Mediterranean,
Iran, and Afghanistan, and its apparent descendants H. dietrichi, H. campbelli, and H.
antelopinum are found in Greece, Iran, and Pakistan, respectively. No certain
members of Hipparion .!•.!• are known from elsewhere, namely East Asia. The reader
should compare Qiu et al.'s (1987) attribution of species to Hipparion (Hipparion) .!•.!•
with Bemor et al. (in press).
Another facet of this issue derives from a recent phylogenetic analysis of North
American Miocene merychippine-grade horses that has been made by Hulbert and
MacFadden (in press). In their analysis, they have generated several alternative
cladograms. A consistent feature of all these cladograms is that Hipparion shirleyi is
the sister taxon of Merychippus insignis, whereas one step higher in their cladograms,
Merychippus insignis is the sister taxon of "Merychippus" (= our Cormohipparion)
goorisi. If one accepts Hipparion shirleyi as the stem North American "Hipparion" .!·.!·'
then this clade is well separated from the Cormohipparion goorisi - Cormohipparion
sphenodus ·- Cormohipparion occidentale - "Hipparion" primigenium - "Hipparion"
~ - Hipparion prostylum transformation series postulated here as the second
phylogenetic hypothesis. Under that scenario, North American Hipparion .!·.!· is a
distinct clade convergent with Old World Hipparion .!•.!• Certainly, this issue is not
currently resolved and is in need of further analysis.
The •stvalhippus Complex•
The "Sivalhippus Complex" includes a number of late Miocene, Pliocene, and

Pleistocene horses ranging throughout Eurasia and Africa. We hesitate to recognize a
302
formal taxon at this time because of the need to study all relevant material, and
resolve the taxonomic muddle surrounding these horses. Indeed, the genotypic species
"Sivalhippus" theobaldi is based only on a juvenile palate fragment from Indo-Pakistan
(GSI C153, from Keypar Punjab, Middle Siwaliks; MacFadden and Woodburne, 1982.;
Bernor and Hussain, 1985), which itself is referred to indirectly associated skull and
postcranial material. Moreover, there are various opinions concerning whether all of
the taxa we review here under this nomen belong to a single genus, or multiple
genera. Despite these qualifications, these horses are united by a number of skull,
mandibular, and postcranial features, including evolutionary trend toward large size,
loss of preorbital fossa, nasals either conservative and undergo no retraction or (as in
Proboscidipparion) they become strongly retracted, cheek teeth become quite high
crowned in this complex, fossette enamel ornamentation stays complex or becomes
even more complex, pli caballins are usually double or multiple, protocones become
triangular to very elongate and triangular, hypoglyphs remain deeply incised, lower
cheek teeth may develop heavily built, columnar protostylids, metaconid and meta-
stylid develop strongly angular facing borders separated by broad, deep U-shaped
linguaflexids, ectostylids may become persistent in more evolved forms, metapodials
tend toward great size, both in length and width, reflecting the generally great size of
the species belonging to this group. Indeed, some species of this group are among the
largest hipparionines known, and represent a poorly appreciated major late radiation
of Old World hipparionines.
Apparent members of the Asian and East African "Sivalhippus Complex" (figure
7) include ".§_." platyodus (ca. 9.5-5 Ma; Bernor et al., in press); ".§_." ptychodus (ca.
8-4.5 Ma; Bernor et al, 1987b, in press; Qiu et al., 1987); ".§_." houfenense (ca. 6-2..5 Ma;
Flynn and Bernor, 1987; Qiu et al., 1987; Bernor et al., in press; MacFadden, 1984); f.·
pater (ca. 5-2. Ma; Qiu et al., 1987); R_. sinense (ca. 3-2. Ma; Qiu et al., 1987; Bernor et
al., in press); ".§.." perimense (ca. 8 Ma; MacFadden and Woodburne, 1982.; Bernor and
Hussain, 1985); ".§_." turkanense (ca. 5 Ma; Hooijer and Maglio, 1976; Bernor and
Hussain, 1985); "S." afarense (ca. 3 Ma; Eisenmann, 1976); "S." rocinantis rocinantis
and".§.." rocinantis crusafonti (Alberdi, 1972., 1974; Qiu et al., l987).
"Sivalhippus" platyodus would appear to be the most primitive member of this

group. It is a medium sized hipparion, with a facial morphology little derived from
"Hipparion" primigenium .!•.!• The relationships of the lacrimal to the POF (character
1c) and absence of a lacrimal foramen (2.b) is duplicated in "H." primigenium s.s. POF
orientation is elongated anteroventrally (3b), its posterior pocketing is reduced (4b/c),
and medial depth (Sb) as well as the peripheral border outline (7b) is relatively
reduced. The nasal notch remains unretracted as in "H." primigenium (14c) and the
maxillary cheek teeth retain complex plications of the pre- and postfossettes (18a),
double pli caballins (2.0a), deeply incised hypoglyphs (Z1b) and lingually flattened and
labially rounded protocones (Z2.b). The lower cheek teeth show some features typical
of this group, namely the molar metaconids and metastylids have sharp angular
opposing borders (30c, 31c, 3Zc, 33c), protostylids are well developed (37a), at least in
the type specimen, but are not yet columnar, and more broadly developed nearly
U-shaped linguaflexids (40d). Other features of the mandibular dentition are conser-
vative and generally "H." primigenium-like. Therefore, it is the reduction of the
preorbital fossa (which is a character found in many independent lineages), the angular
morphology of molar metaconids and metastylids, deep U-shaped linguaflexids in
molars, and the apparent lack of any autapomorphies that mark this as potentially the
most primitive known derivative of the "Sivalhippus Complex."
"Sivalhippus" ptychodus is very similar in size and morphology to ".§.." platyodus;

it is derived from the same area in China (Wuxiang) and was synonymized with the
latter taxon by Qiu et al. (1987). However, Bernor et al. (in press) have cited some
consistently divergent characters typical for this group. Namely, the preorbital fossa
is somewhat further reduced and placed further anteriorly on the face with respect to
the lacrimal (character 1g). The preorbital fossa is smaller (3j), similarly pocketed
(4b), but medial depth is decidedly greater (Sa). The nasal notch is not well preserved
on any specimens, but does not appear to be retracted beyond PZ/ (14?c). The maxil-
lary cheek teeth have very complex plications of the pre- and postfossettes (18a+/a),
303
Ma EUROPE C H I N A AFPICA
2
~
~~~.::,)l?l") ~ ::""i--
,-~·~.
3 ~ ' • - ~
' ~ P si nensis ~!/S. afi.houlenense ""S:" afO<ense
1
' S:•crusalonll ~" J
··s:- r, roc:lnentls
p :-~ ~
I
~ "S~ turkanense
~S7 houfenense
-~
"s:- ptyehodus
8
·~ ..$ :" perlmense
10 ~ ~otyodus
11 ~H~ Drlmlae~lum s.s..
Fig. 7. Evolutionary history of the "Sivalhippus" Group. From the lower left
comer, a species close in morphology to "Hipparion" primigenium s._!.
evolves to the "Sivalhippus" platyodus state; "~·" platyodus is the
potential sister-taxon of "~." ptychodus (Baodean-Gaozhuangian) in
China, and "~·" perimense species (L. Miocene) in Indo-Pakistan,
respectively; "~." perimense is the sister-taxon of the Chinese taxon,
"S." houfenense (Baodean, Gaozhuangian, and Youhean); "S."
houfenense is potentially related to the ProboscidippariOn
(?= "Sivalhippus") pater (Gaozhuangian-Youhean), Proboscidipparion
(?= "Sivalhippus") sinense and "Sivalhippus" aff. houfenense (Youhean)
clades, respectively; "Sivalhippus" perimense also is potentially related
to the East African transformation series ·~." aff. perimense - "~."
turkanense - "S." afarense. "S." houfenense would also appear to be
related to the Spanish species "s." rocinantis (Ruscinian-Villafranchian
age). -
pli caballins are double or complex (ZOa/c). All other features of the maxillary cheek
teeth are duplicated in "H." platyodus. Unfortunately there are no certainly associ-
ated mandibular or postcranial remains to evaluate.
Another Asian late Miocene representative of this group is "Sivalhippus"

perimense (Bemor and Hussain, 1985; = "Hipparion" feddeni, in part, of MacFadden and
Woodburne, 1982). There is considerable variability in measurements given for this
taxon, and indeed more than one species may be included here (Bemor and Hussain,
1985). The best represented collection is at the American Museum, collected by
Barnum Brown, and is believed to originate from the 8 million year level of the Potwar
Plateau (Barry, personal communication to Bemor). It is this sample that we specifi-
cally refer to in our characterization.
This is a large horse, with preorbital fossa relatively reduced in size, with a long
preorbital bar placed far anterior to the lacrimal bone (character 1g). The fossa is
approximately egg-shaped and anteroposteriorly directed (3d), has no posterior pocket-
ing (4c), retains moderate depth (5b), strong peripheral border outline (7a), and a
304
distinct anterior rim (Sa). The nasal notch remains unretracted, being positioned at
the anterior border of P2/ (14c). The maxillary cheek tooth occlusal morphology is
mostly conservative, retaining complex plications of the pre- and postfossettes (18a),
double pli caballins (20a), and deeply incisea hypoglyphs (21 b). However, cheek tooth
mesostyle crown height is great, 60+ mm in unworn crown height (17d), and protocones
are distinctive of this group in having acquired a more triangular shaped morphology
(character 22e). The lower cheek teeth show a number of derived features of this
group including strongly angular facing borders of metaconid and metastylid (30c, 31c,
32c, and 33c), strongly built, columnar protostylids (37c), and deep U-shaped lingua-
flexids (39d, 40d) which accommodated occlusion with the expanded triangular-shaped
protocones (Bernor, personal observation of AMNH collection). Undirectly associated
postcranials in the AMNH are elongate and robustly built (Bernor and Hussain, 1985).
Qiu et. al. (1987) have recently reported on a suite of fine specimens of
"Sivalhippus" houfenense. This horse would appear to have first occurred in the latest
Miocene, ca. 6 Ma, and persisted well into the Pliocene, ca. 2.5 Ma. It overlaps con-
siderably in size with "H." primigenium and ".§_." perimense, but no ".§.." houfenense
individuals are as large as the largest members of these other taxa. The preorbital
fossa is sharply reduced in all its dimensions and features, and as an apparent conse-
quence of this, the preorbital bar appears to have been somewhat shortened (Qiu et
al., 1987; Table 1, pg. 36, measurement of ca. 32 mm seems very short for this line-
age). The maxillary cheek teeth retain largely primitive features including complex
fossette plications (18a), mostly double pli caballins (20a), and deep hypoglyphs (21 b);
however, protocones are derived in their triangular morphology (22e). The mandibular
dentition likewise reveals a number of derived features including incisors that are
grooved with strongly developed lingual pillars (27b), while remaining curved (28a) and
with I/3 not transversely constricted (29a); cheek tooth metaconids and metastylids
have sharp angular facing borders (30c, 31c, 32c, 33c); protostylids (37a) are present,
but not so strongly built; ectostylids are not consistently present (38b); linguaflexids
are broadly U-shaped (39d, 40d).
A larger and later specimen, F:AM 111820 was referred to "Neohipparion"

houfenense by MacFadden (1984; with some hesitancy on his part) and later by Flynn
and Bernor (1987) to "Hipparion" aff. houfenense. This is a somewhat larger horse
than "S." houfenense s.s., and it is known from later Pliocene localities that also
contaiii"Equus sanmanensis. This taxon is derived in its complete loss of a facial fossa
(characters lh, 2b, 3g, 4e, Sd, 6a, 7e, 8b). The nasal notch is plesiomorphic in its
retraction only to the anterior surface of P2/ (14c). The cheek teeth have somewhat
reduced fossette plication complexity (18b), but pli caballins remain double (20a) and
hypoglyphs are deeply incised (21 b). Protocones are derived in their triangular
morphology (22e). Likewise, the lower dentition shows the derived pattern of incisor
morphology (27b; but 28a and 29a), metaconid, metastylid, and linguaflexid (30c, 31c,
32c, 33c and 39d, 40d) are typical for this group, and with protostylids entirely lacking
(37b).
"Sivalhippus" houfenense-like taxa would also appear to be represented in

European Pliocene aged horizons. Alberdi (1972: Figs. 96-99) has figured specimens of
two hipparions which she refers to H. rocinantis rocinantis and H. rocinantis
crusafonti, arguing that they are time-successive chronosubspecies. We have not
studied these specimens, but the figures and descriptions are revealing of this species'
phylogenetic relationships. We follow Zhegallo (1978) and Qiu et al. (1987) in recog-
nizing the strong apparent resemblance between "S." rocinantis and "S." houfenense
(sensu lato), as well as African members of this clade. The Skull fragment
(MNCM 2268, from the Villafranchian locality of Villaroya) is virtually identical also
to AMNH 111820 in preserved morphological features of the facial region and maxil-
lary cheek teeth (except 18a; some apparently more complex plication). Mandibular
cheek teeth of "S." rocinantis crusafonti (IPP-Sabadell No. V.175) are also virtually
identical to the AMNH specimen. Metapodials are long and robustly built.
Africa likewise would appear to harbor representatives of this clade, including

the latest Miocene taxon "S." turkanense (Hooijer and Maglio, 1974) and the Pliocene
305
taxon "~_." afarense (Eisenmann, 1976). Eisenmann (198Z), Bernor and Hussain (1985),
and Bernor et al. (1987b) have remarked that ".§_." turkanense is remarkably similar to
the Siwalik horse ".§_." perimense in all morphological features of the skull. In compar-
ison, however, the preorbital fossa is further reduced (3?g, 4d, Sc, 7d, 8b), with only a
remnant depression placed far anterior to the anterior tip of the lacrimal bone. The
maxillary cheek teeth likewise retain all the essential features of the Siwalik
species. Hooijer and Maglio (1974) referred some lower cheek tooth specimens from
not only Lothagam (Pl. 7: Figs. 1-Z, KNM LT-171; Figs. 3-4, KNM LT-170) and Ekora
(Pl. 7: Figs. 5-6), plus Hipparion primigenium from Lothagam (Pl. 6: Figs. 3-4) and
Kanapoi (Pl. 6: Figs. 1-Z), that share characters with species of the "Sivalhippus
Complex."
"Sivalhippus" afarensis is very similar to ".§_." turkanense in its facial and maxil-
lary morphology. Eisenmann (1976) has reported an associated partial skull and com-
plete mandible from the Afar triangle, circa 3.6 to 3.0 Ma (compare Boaz et al., 198Z
and White et al., 1984 for arguments about the chronology). This is a very interesting
specimen, and potentially important for phylogenetic linking of late Miocene and Plio-
Pleistocene taxa belonging to this clade. This is an early to middle stage-of-wear
adult, with M3/'s occlusal surface well worn, and PZ/ mesostyle height approximately
46 mm. The preorbital bar is very long (54.9 mm) with the anterior edge of the
lacrimal placed less than one-half the distance from the anterior orbital rim to the
posterior rim of the fossa (lg). The preobital fossa is vestigial, but distinct (3g), has a
distinct posterior rim (4c), medial depth is quite shallow but discernible (Sc), and
peripheral outline is virtually indistinct (7d). Other features of the facial region are
as in members of this clade described above (Zb, 6a, 8b, 9b, lOb, lZa, 13a, ?14c).
Maxillary cheek teeth are also similar to members of this group (18a, 19b, ZOa, Zlb),
but protocones show the triangular morphology to some extent on P4/-MZ/, while
being more oval-elongate in shape on the other cheek teeth. This may be wear
related, or mark a morphological transformation in this lineage. As in more derived
Asian species of this clade, "S." afarense has mandibular incisors which are grooved
(character Z7b) and markedly straight (Z8b); cheek tooth metaconids and metastylids
have angular facing borders (30c, 3lc, 3Zc, 33c), ectoflexids are quite shallow on the
premolars (34a), while they separate metaconid/metastylid on the molars (35b); pli
caballinids are complex to single (36a/b); protostylids are small when present (37b);
ectostylids are expressed on P/3, M/1, and M/Z only (38a); linguaflexids are deep and
very broad (39d, 40d).
Eisenmann (1979) has made some further important observations on Plio-Pleisto-

cene hipparionines. A suite of dental characters appear during this interval (ca.
3.6-Z.S Ma): (1) reduction of 13/3 (especially marked in I/3 (character Z9b); (Z) appear-
ance of ectostylids in adult dentitions (character 38a). Later in the sequence (less
than Z Ma), mandibular symphyses become very broad with I/1-Z being hypertrophied,
and more anteriorly directed (Z8b) with accompanying sharp reduction of I/3 placed
immediately posterior to I/Z. These characteristics are seen in a specimen from
Olduvai Gorge locality BK ll (Old SZ 067/5344; cast in MNHN). Churcher and
Richardson (1978) have referred the youngest African material, containing all the
apomorphies cited above, to three subspecies of Hipparion (= Stylohipparion of earlier
usage) libycum: H. libycum libycum in North Africa, H. libycum ethiopicum in East
Africa, and H. libycum cornelicum in South Africa. A closer analysis of these "super-
specific" relationships could potentially be quite revealing about this highly apomor-
phic lineage's late evolution and potential biogeographic extension throughout Africa.
Two Plio-Pleistocene Asian species of Proboscidipparion, ~· pater and P. sinense,

share a number of the cheek tooth and postcranial features cited above for the
"Sivalhippus Complex." However, they differ significantly in their very strong retrac-
tion of the nasals. It is principally for this reason that Qiu et al. (1987) and Bernor et
al. (in press) have at least provisionally retained the generic nomen Proboscidipparion.
Qiu et al. (1987) have recently reported on the Pliocene species

Proboscidipparion pater (ca. 5-Z Ma). This species completely lacks a preorbital fossa
(characters lh, Zb, 3g, 4e, Sd, 6a, 7e, 8b). The nasals are strongly retracted to a
306
position of M1/ mesostyle (14g). The cheek teeth have very richly plicated fossette
margins (18a+), pli caballins are double (20a), hypoglyphs are deeply incised (21b).
Protocones are conservative, being lingually flattened and labially rounded (22b).
Mandibular incisors would not appear to be grooved or to have lingual pillars (27a), and
are curved (28a). I/3 is not transversely constricted (29a), premolar and molar meta-
conids and metastylids have angular shaped facing borders (30c, 31c, 32c, 33c), and
linguaflexids are deep U-shaped structures (39d, 40d). It is not clear if the mandibular
cheek teeth have distinguishable protostylids, but ectostylids seem to be absent
(38b). The metapodials are elongate and robust.
Proboscidipparion sinense is approximately 1/7 larger than !:· pater and first
occurs in medial Pliocene times (ca. 3 Ma; Qiu et al., 1987; Bernor et al., in press).
Proboscidipparion sinense has no facial fossa (characters 1-8 as for!:· pater), but the
nasals are retracted to a position nearly over the orbits (character 14h). The cheek
teeth have very complex fossette plications (18a+), pli caballins are remarkable for
their great complexity (ZOe), hypoglyphs are deeply incised (21b), and protocones have
a derived triangular-elongate shape (22f). The mandibular dentition is quite similar to
!:· pater, except for the very derived condition of canine loss, and the very long and
relatively narrow dimensions of the cheek teeth as a whole. Protostylids are small and
impersistent in this species (37b). The metapodials are very long (perhaps the longest
of all hipparions), and robust.
The taxa we have brought together into the "Sivalhippus Complex" constitute a
radicle of late Miocene-Pleistocene aged hipparions, united by a number of skull,
mandibular, and postcranial characteristics. The precise phylogenetic relationships of
this complex, whether its diversity includes a single genus or multiple genera (i.e., the
potential validity of the genera Sivalhippus, Proboscidipparion, and Stylohipparion),
must yet be evaluated.
Cremohipparion
Woodburne and Bernor (1980) recognized a suite of eastern European, western

U.S.S.R., and southwest Asian hipparions characterized principally as having a short
preorbital bar with lacrimal bone invading the posterior wall of the fossa, and termed
these Group 2 horses. Subsequent work by Bernor et al. (1980), Bernor and Hussain
(1985), Bernor (1985a), and Bernor et al. (1987b) presented geochronologic aspects of
these taxa. Qiu et al. (1987) have made a very important contribution in not only
recognizing species of these horses in China, but erecting the subgenus Hipparion
(Cremohipparion). We believe that the suite of species we cite below as belonging to
this taxon constitute a natural group, and are a lineage quite separate from Hipparion
.!·.!· in the New or Old Worlds, and also are distinct from Group 1 taxa or the
"Sivalhippus Complex." Bernor and Tobien (1989) have raised Cremohipparion to full
generic rank.
Species which we include in Cremohipparion include (figure 8) C. moldavicum

(ca. 9-8 Ma; Gromova, 1952; Woodburne and Bernor, 1980, Fig. 3F but not Fig. 6H, and
relevant text, p. 1131, 1342-43; Bernor, 1985a); C. matthewi (Woodburne and Bernor,
1980; Bernor, 1985a); C. mediterraneum (ca. SMa; Woodburne and Bernor, 1980;
Bernor, 1985a); C. proboscideum (ca. 8-7 Ma; Sondaar, 1974; Woodburne and Bernor,
1980; Bernor, 1985a); C. aff. mediterraneum (late Miocene; Bernor, personal observa-
tion); C. forstenae (ca. 9-7.5 Ma; Bernor et al., 1987b, in press; Qiu et al., 1987); C.
licenti (ca. 5+-3 Ma; Bernor et al., 1987b, in press; Qiu et al., 1987). Other apparent
members of this group include "H." tudorovense (Gabunja, 1959, Pl. V, Figs. 1,2), "H."
gromovae (Gabunja, 1959, Pl. ll, Figs. 1-3), "H." sp. (Gabunja, 1959, Pl. Vll, Fig. 1).
Cremohipparion moldavicum is known from a series of localities in the western

U.S.S.R. (Gromova, 1952; Gabunja, 1959), and specimens which are referable to this
taxon have been reported from Maragheh, Iran (Bernor, 1985a). This species is
remarkable for being only medium size, but retaining a large preorbital fossa with a
lacrimal that invades its posterior border (character 1b). There is no lacrimal foramen
(Zb). The preorbital fossa is subtriangular shaped and anteroposteriorly oriented (3c),
posterior pocketing is sharply reduced and slight (4b). Medially, the fossa is deep (Sa),
307
,. E · MEDITERRANEAN, W. USSR. S..W. ASIA CH I N A
C. llce-ntJ
~\
6
~-. C. probo•cldeum \ . : .
C. forstenee
8
~\~' _....,.-~
C. matlhewJ -
~
C. aU. medlterraneum
C. mediferraneum
C. moldovic:um
10 ~ u P e
-~
11 ~
"'H ~ pr lmlge-nlum
12
C. .sphenodus
13
Fig. 8. Evolutionary history of the Cremohipparion Group. From the lower

right, Ho: 1 (Hypothesis 1, of Woodburne et al., 1980), the origin of
Cremohipparion from the North American Cormohipparion sphenodus
stage-of-evolution; from the lower left, Ho: Z (Hypothesis Z, of
Bernor, 1985a; Bernor and Hussain, 1985; and Bernor et al., 1987), the
origin of Cremohipparion from the "Hipparion" primigenium s.s. stage-
of-evolution; in the western U.S.S.R., eastern Mediterranean, south-
west Asian region, Cremohipparion moldavicum (Turolian = Meotian)
differentiates in the early Turolian from central European "Hipparion"
primigenium .!•.!•' followed by the differentiation of the C.
mediterraneum (E. Turolian) - C. proboscideum (M. Turolian) and C.
matthewi (M. Turolian) clades; in China, there is evidence for the
occurrence of C. aff. mediterraneum (Baodean) from the late Miocene
of Shouyang (Bernor and Qiu, in progress), which is a plausible sister
taxon of "C." forstenae (Baodean) and "C." licenti (Gaozhuangian).
lacks internal pits (6a), has a strong peripheral border outline (7a), and anterior rim is
distinct (Sa). The infraorbital foramen is placed inferior to, and encroaches upon, the
anteroventral border of the preorbital fossa (9b). The buccinator fossa is distinct
(lOb), but not posteriorly pocketed (lla), and there is no caninus fossa (lZa). There is
no malar fossa (13a). The nasal notch is incised only to the anterior border of PZ/
(14c). There is no persistent and functional Pl/ (15b). The maxillary cheek teeth are
moderately curved to straight (16b) and maximum crown height is between 40 and
60 mm (17 c). Maxillary cheek tooth fossette ornamentation is moderately complex
(18b); posterior wall of the postfossettes are always distinct (19b); pli caballins are
variably single or double (ZOa/b); hypoglyph is moderately deeply incised (Zlc); proto-
cone is rounded (ZZd), clearly isolated from the protoloph (Z3b), has no noticeable
protoconal spur (Z4c), and is lingually placed relative to the hypocone (ZSb). PZ
anterostyle/paraconid are elongate (Z6a). Mandibular incisors are not grooved (Z7a),
they are curved (Z8a), and I/3's are not transversely constricted (Z9a); premolar and
molar metaconids and metastylids are rounded (30a, 3la, 3Za, 33a); ectoflexids do not
separate metaconids and metastylids on the premolars (34a), but they do on the molars
(35b); pli caballinids are generally absent (36c); protostylids are very rare (37b), ecto-
308
stylids absent (38b), and linguaflexids are shallow to shallow V-shaped (39a/b), whereas
in the molars they are somewhat deeper and V-shaped (40b).
Cremohipparion matthewi was provisionally referred by Woodburne and Bernor

(1980) to Group 4. Bernor (1985a) referred "Hipparion" matthewi to Group Z of
Woodburne and Bernor (1980) based on a Maragheh specimen (GIU P100-1958) which
seemed to conform to published descriptions and poor photographs of the type speci-
men. At the invitation of L. Kordos, Bernor and Tobien (1989) recently studied the
type specimen, and verified that it indeed belongs to the Cremohipparion lineage. C.
matthewi proves to be quite like C. moldavicum in many morphological features, and
indeed is generally found in the same geographic region, namely Greece, Iran, and the
western U.S.S.R. It shares with C. moldavicum the plesiomorphic feature of lacking a
distinct caninus fossa (character1Za). It differs from C. moldavicum in its consis-
tently smaller, dwarf size and its primitive retention ofan anteroventrally oriented
preorbital fossa (3b). The preorbital fossa is diminished in its expression with reduced
posterior pocketing (4c), medial depth (5c), peripheral border outline (7b), and loss of
an anterior rim (8b). The type differs from the Maragheh specimens in cheek tooth
morphology by more consistently retaining double pli caballins (ZOa; versus ZOb in the
GIU P100-1958 specimen). Unlike C. moldavicum, protocones show more variability
between the primitive lingually flattened-labially rounded (ZZb) and rounded states
(ZZd). Mandibular specimens are somewhat better known for C. matthewi and differ
from "C." moldavicum only in having a distinct gradient in linguaflexid morphology,
being generally shallow in the premolar series (39a), and morphologically deeper and
U-shaped in the molar series (40c).
C. mediterraneum, and indeed all of the remaining hipparions listed here, are
derived compared to C. moldavicum and C. matthewi in having a distinct, well
defined, caninus fossa (character 1Zb). C. mediterraneum also differs from C.
moldavicum in having an anteroventrally oriented fossa (character 3b), the nasals are
retracted to a position above the mesostyle of PZ/ (14d), fossette plication is complex
(18a), pli caballins are consistently double (ZOa), hypoglyphs are deeply incised (21 b),
and protocone shape varies from being lingually flat and labially rounded (ZZb/d). A
specimen from Shouyang, China, housed at the AMNH, and under study by Bernor and
Qiu, is virtually identical to C. mediterraneum in size and every morphological
attribute.
C. proboscideum from Samos Greece (Wehrli, 1941; Sondaar, 1974; Woodburne

and Bernor, 1980) is a larger horse than those described above, and remarkable for its
elongate snout, hypertrophied preorbital fossa, and more strongly retracted nasals to
mesostyle of P3/ (character 14e). Other than these features, there is little to distin-
guish it from C. mediterraneum.
C. forstenae is a Chinese horse, particularly well represented from the Baode

County, Shanxi Province (Sefve, 1927; Bernor et al., 1987b; Qiu et al., 1987). It is
derived compared to C. mediterraneum in its reduced expression of the preorbital
fossa, namely lack of posterior pocketing (character 4c), reduced medial depth (5c),
and weakly defined peripheral rim (7c). As in C. proboscideum, the nasals are
retracted to a level of P3/ mesostyle (14e). The cheek teeth show some derived char-
acters, including reduced complexity of the fossette margins (18b), pli caballins vari-
ably double or single (ZOa/b), hypoglyph incised variably from moderate to shallow
(Zlc/d), protocone variably oval to rounded (ZZc/d), with a greater propensity toward
the latter state in middle adult wear.
C. licenti is chronologically the youngest and morphologically the most derived

member of the group. It is unique among all hipparionines in the occurrence of inter-
nal circular pits within the preorbital fossa's medial wall (character 6b), a posteriorly
pocketed buccinator fossa (llb), and a distinct malar fossa (13b). Other derived
characters, compared specifically to other species of Cremohipparion listed above,
include the strongest nasal retraction (to mesostyle of Ml/; 14g), extreme simplicity
of fossette plication frequency and amplitude (18c), consistently single pli caballins
(ZOb), shallowly incised hypoglyphs (Z1d), and rounded protocones (ZZd).
309
Two distinct hypotheses of this lineage's origin have been advanced. Woodburne
et al. (1981) proposed Cormohipparion sphenodus, from the late Barstovian and early
Clarendonian of North America (ca. 14-12 m.y.), as the most likely ancestor, and its
first occurrence in the Old World represented a second, later (Turolian) immigration.
Woodburne et al. (1981) argued that Group 1 and Group 2 (= Cremohipparion here)
horses must have separated phylogenetically by the early Vallesian, and an unknown
common ancestor must have had a relatively derived (in comparison with North
American Cormohipparion) subtriangular shaped and anteroventrally oriented POF.
Woodburne et al. (1981) further postulated that Group 1 became larger, increased
dental complexity, attained a longer preorbital bar, and diminished strength of the
anterior rim. Following their interpretation, Group 2 horses retained a relatively
smaller size and reduced preorbital bar length so that the lacrimal contacted the
posterior border of the preorbital fossa. Woodburne et al. (1981) believed it unlikely
that Group 2 horses secondarily reduced the preorbital bar from a Group 1 ancestor.
They advanced the hypothesis that Cormohipparion sphenodus represents a plausible
sister taxon for both Old World Group 1 and Group 2 forms, in contrast to Cormo-
hipparion occidentale.
In contrast to that scenario, Bernor and Hussain (1985), Bernor (1985a), and
Bernor et al. (1987b) advanced the hypothesis that Cremohipparion was derived from
"H." primigenium, or a close relative, which in turn was morphologically and hence
phylogenetically very closely related to Cormohipparion occidentale (see arguments
supporting this assertion above).
An evaluation of character state distribution here allows us to illustrate the

problem. Characters which clearly support Woodburne et al.'s (1981) hypothesis
include the lacrimal's contact with the posterior wall of the preorbital fossa (char-
acter 1b) and concomitantly narrow preorbital bar in both Cormohipparion sphenodus
and all Cremohipparion species. However, a number of characters support a phylo-
genetic relationship between "H." primigenium s.s. and Cremohipparion, including the
lack of a lacrimal foramen (character 2b), presence of an anteroventrally oriented
preorbital fossa in at least some species belonging to the group (3b), nasals retracted
closer to P2/ (14c) or posterior to P2/ (14d, e, g), P1/ generally absent (15b), presence
of complex plications in some forms of Cremohipparion (i.e., C. mediterraneum and C.
proboscideum; 18a). Also, it should be noted that one species of Cremohipparion, C.
proboscideum rivals "H." primigenium .!·.!· in size (perhaps a secondary acquisition
within the Cremohipparion clade) while the remaining and geologically earlier taxa are
smaller. There are a number of characters shared by both Cormohipparion sphenodus
and "Hipparion" primigenium s.s. with Cremohipparion (i.e., 3, 7, 8, 19, 21, 25, and all
mandibular characters), which for the most part appear to be symplesiomorphies.
Chief characters which support the proposed Cormohipparion sphenodus-Cremo-
hipparion relationship (contact of the lacrimal with the preorbital fossa and short
preorbital bar) might be alternatively explained as reversals.
Postcranial Evidence
In table 2 we have presented all available measurements of North American and

Old World hipparionine MC m•s and MT m•s. We summarize here some patterns of
these limb bone proportions after bivariate plots of maximum length (var. 1) versus
distal articular width (var. 11). As in figure 4, we use the HOwenegg 95% and 99%
probability ellipses as a standard for population comparison. We wish to qualify that
most of the localities we refer to here, have two or more hipparion species, and
referrals of postcranial elements to species, which are mostly based on skulls and
mandibles, is not absolutely certain. We have scrutinized these data carefully, and
offer our best judgments here.
Figure 9 depicts the MC m proportions: 9A gives all available Old World

hipparionine measurements together in a single plot, while 9B through 9E segregate
these measurements by the different Old World evolutionary groups that we have
defined on the basis of skulls, mandibles and dentitions. The numbers of the different
taxa refer directly to those given in tables 1 and z. Figure 9B, Old World "Group 1"
310
Mltacarpals III MltoCII'IIIll Ill
350 3liO I
A B
300 3110
,.16
j 250 .,25 ,;Ill 1 250
~. ~
I
:;
o3B
16~
37;• ~.il.o.l7 I 200
1&,~-
200
i 23 5-:li'
~
i
150 150
100 I
100 .
15 25 35 <15 !1!1 15 25 35 <15 !15
DAif lool DAif lool
Mltocorpols III Mltocorpsls Ill
350 350 I
I I
c D
300 300
,.16
1
~
250 o25'
e2ol
1
:;
250
... ~
,;Ill
!
i
200 o23ce;P
I
i
200 ct:::P
~
150 150
I I I I
100 100
15 25 35 <15 55 15 211 3!i e !15
DAif lool DAif lool
Mltocorpolo III
350
E
300
] 250 ~
I 200 o3B "rl~

i
150
I
100
15 211 311 «< !1!1
D.IW Ill)
Fig. 9. Bivariate plots comparing Old World hipparionine MC m maximum l~ngth

(var. 1) versus distal articular width (var. 11) proportions with 95% and 99%
Howt;!negg confidence interval ellipses superimposed. A. All available
species. B. "Group 1." C. Hii!J!arion s.s. D. "Sivalhii!J!US Complex."
E. Cremohii!J!arion. Note that the num btirs refer to those taxa listed in
tables 1 and z.
311
hipparions, reveals that most species fall within the 95% and 99% confidence limits of
the Howenegg sample (note that "H." primigenium s.s. is represented by four distinct
populations: lZa- Eppelsheim, lZb- Howenegg, lZc- Charmoille, lZd- Vienna
Basin). The exceptions are numbers 16, "H." melendezi and 22, "H." hippidiodus
(although these are both found relatively close to the others). As a "group" then, these
species show very little to no divergence in their proportions away from the Howenegg
horse sample. Figure 9C, Hipparion s.s., is represented here by H. prostylum (23), H.
dietrichi (24), and H. campbelli (25). All these species fall outside of the Howenegg
ellipses, and interestingly the most primitive member of Hipparion s.s., H. prostylum,
falls closest to the regression of the Howenegg horse, having therefore relatively
similar length to width proportions. However, H. dietrichi (Z4) and H. campbelli (Z5)
are progressively more advanced in their greater length to width proportions. Figure
9D, the "Sivalhippus Complex," again has all representative species with MC m
measurements 1 X 11 falling outside the Howenegg 95% and 99% ellipses. Here,
however, there is an intragroup diversity in limb proportions. "S." houfenense (30),
".§_." rocinantis crusafonti (32), and Proboscidipparion pater (35) would appear to be on
a common intragroup regression line that parallels the Howenegg sample, and has
larger size reflected concommitantly by their greater length and width. In contrast,
".§.." platyodus (Z7) has marked length reduction of MC m and shows a tendency for
dwarfing. Quite the contrary, Proboscidipparion sinense (36) is divergent from all
other members of this group in its extremely elongate MC m proportions. Figure 9E,
the Cremohipparion species, C. moldavicum (37), C. matthewi (38), C. mediterraneum
(39), and C. proboscideum (40) also show some morphological diversity in their limb
proportions. C. "matthewi" is a dwarf form with a relatively very similar slender
MC ill; indeed, there is more than one dwarf species of Cremohipparion in the Samos
fauna, and it is currently impossible to say which species (if not both) of these dwarf
hipparions is associated with the postcranials we figure here (Bernor and Tobien,
1989). C. proboscideum (40) would appear to have relatively long and slender limbs
compared both to "Group 1" species and other members of the Cremohipparion group
presented here; however, since there is such a great diversity of hipparions from
Samos, this postcranial referral is made very tentatively. C. moldavicum (37) and C.
mediterraneum (39) have slightly more slender MC ill proportions than "Group P'
species, but would appear to be more conservative than the other two Cremohipparion
species in their divergence away from the MC ill proportions exhibited by "Group 1"
species.
The MT m data (figure lOA) reflects very much the same patterns as found with
the MC m data with some minor exceptions. In figure lOB, Old World "Group 1"
species find a proportionally shorter length dimension for "H." hippidiodus (16) than
was found for MC m. In figure lOC, Old World Hipparion s.s., there are no complete
metatarsal ill's referable to H. campbelli (25), but H. dietrichi (Z4) shows a sharp
increase in length to width (more gracile) limb proportions compared to H. prostylum
and "Group 1" hipparionines. In figure lOD, the "Sivalhippus Complex" and figure lOE,
the Cremohipparion lineage, our bivariate plots show much the same pattern of length
to width proportions than the MC ill data have.
Bernor et al. (1988: 433, fig. 5) have demonstrated that Central European
Vallesian "Hipparion" primigenium lived in warm temperate mesophytic forests with
little to no winter frost. Tobien (1952) noted that the Hl:iwenegg hipparion foot bones
reveal a likely functional monodactyly during fast, open country locomotion, but
during slow locomotion digits II and IV were fully functional in facilitating movement
over soft terrain. Those species cited above with relatively elevated length to width
(more gracile) proportions may have had adaptations to more cursorial activity,
whereas those with lower length to width (more robust) proportions may have had
adaptations for a slower mode of locomotion (Gromova, 1952; Forsten, 1968).
Discussion
The morphological distinction of several Old World hipparion lineages, and their
apparent North American sister and outgroup taxa, serves as sufficient evidence to
justify ours and previous workers' assertions that hipparions cannot be included in the
312
Metatarsals· III Metatarsals III
350 350
I I
A o36
B
300 300
0 24 e]O, 32
o40 o35
I 250 3 ~'112ci1))
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.
22o 250
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~
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0
rs 12d 12a, b. 13
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200 200
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=
150 150
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CAW (111111) CAW (rnm)

Metatarsals III Metatarsa Is III
I I I
c 0 o36
300 300
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o35
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CAW (mill) DAN 1om)

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300
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··3~
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37
200
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DAN 1..1
Fig. 10. Bivariate plots comparing Old World hipparionines MT ill proportions with
95% and 99% Howenegg confidence interval ellipses super imposed. A. All
available species. B. "Group 1." C. Hipparion ~·~· D. "Sivalhippus
Complex." E. Cremohipparion. Note that the numbers refer to those taxa
listed in tables 1 and 2.
313
single genus Hipparion, The lineages which we have characterized here are united by
distinct morphological characters, and are spatially and temporally bounded. Bernor
et al. (1987b) reviewed the biogeographic relationships of these groups based on an
earlier and less explicit evolutionary survey. We believe that our analysis bears rele-
vant information not only on evolutionary relationships, but also on biogeographic and
chronologie distribution, patterns of evolution, whether cladogenetic or anagenetic,
and generalizations about times of climatic change on a continental to global scale,
with resulting biogeographic connections.
"Group 1" species are the most primitive Old World hipparionines. They are
distributed in Europe, North Africa, West Asia, and western U.S.S.R., North and East
Asia between 11.5 and 5 million years ago. It is probable that related species were the
first to migrate into Indo-Pakistan and Sub-Saharan Africa, albeit somewhat later in
time (ca. 10-9.5 Ma). We have suggested here that the most primitive taxon of this
"group," "Hipparion" primigenium s.s., is the sister-taxon of North American Cormo-
hipparion occidentale (Bernor et al., 1988; also this agrees with an independent phylo-
genetic analysis by Hulbert, personal communication to Bernor), and that more derived
species of this "group" are no more than provincial, conservative, evolutionary
branches. At the same time, work in progress by MOW indicates (as with Eisenmann
et al., 1987) that specimens now included in Cormohipparion occidentale may pertain
to more than one species and that it remains to be determined if any of these species
was the more likely ancestor for "Hipparion" primigenium. Assuming that a species of
Cormohipparion gave rise to "H." primigenium, the pattern of evolution by this "group"
of horses would appear to be cladogenetic by means of geographic vicariance. The
cause of vicariance in most cases would best be explained as being environmental, or
in a more succinct sense, perhaps ecological differentiation accommodated by geo-
graphic distance. Hypotheses of punctuation versus gradualism are untestable insofar
as we cannot evaluate them in a single section of rock for this "group," or any other
hipparionine lineage (sensu Dingus and Sadler, 1982), and therefore are superfluous to
our argument here.
Old World Hipparion !•!• had a geographic range extending from western Europe
to the eastern Mediterranean, southwest Asia, and Indo-Pakistan, where it is relatively
scarce. The origins of this group are still controversial, yet Old World members reveal
some distinct evolutionary and biogeographic patterns. Hipparion prostylum is recog-
nized as being the earliest Old World member of this group. Bernor (1985a) and Bernor
et al. (1987b) have suggested that H. prostylum extended the range of this lineage to
its maximum Old World geographic extent, and then, due to an environmentally
induced vicariance (circa 8 Ma), species populations were geographically fragmented.
The result was the emergence of a diverse set of provincial autapomorphic species: H.
dietrichi in Greece, H. campbelli in Iran, and H. antelopinum in Indo-Pakistan. This
pattern would again be a cladogenetic one, most probably induced by environmental
vicariance. However, Bernor (1985a) cites a potentially direct, and suitably strati-
graphically continuous anagenetic evolution between Middle Maragheh H. prostylum
and H. campbelli, revealing a different, local mode of evolution (see also Woodburne
et al., 1981).
The "Sivalhippus Complex" includes a large suite of very diverse hipparions

ranging in age from 8+ to about 1 Ma, and distributed pr marily in Asia and Sub-
Saharan Africa, but with at least some Peri-Mediterranean representatives also.
There would appear to be two intervals of biogeographic extension: first, circa 8 Ma
between Indo-Pakistan and Sub-Saharan Africa, with potentially isolated evolution in
Africa thereafter; then later, circa 5 Ma, into western Europe and China. Whereas the
European extension does not appear to have resulted in a remarkable cladogenesis of
this group, the Chinese radiation did, with the Proboscidipparion and "Sivalhippus"
clades apparently diverging during the early Pliocene.
The Cremohipparion lineage was distributed in the western U.S.S.R., eastern

Mediterranean, southwest Asia, and east Asia between circa 9 and 3 Ma. The earliest
and most primitive species of this group appears to have occurred in the eastern
Paratethys and Iranian regions (i.e., Peri-Caspian Sea area of today) and possibly were
314
derived from a species not morphologically dissimilar from "Hipparion" primigenium
s.s. (or its immediate ancestor). This group appears to have extended its range into
Asia circa 8+ Ma, and subsequently fragmented into a western (eastern Mediterranean-
southwest Asian) and eastern (Chinese) radicles. This event would have most probably
occurred between 8 and 6 Ma, but remains to be clarified by ongoing geochronologic
work in China (Qiu, Tedford, Flynn, and others, in progress). The pattern of biogeo-
graphic extension and subsequent vicariant diversification is similar to that cited
above for the other Old World hipparionine lineages.
There appears to have been a number of distinct Old World hipparionine evolu-
tionary radiations. The first was a period of initial differentiation of "Group 1"
hipparions which occurred almost immediately after the first appearance of the
"group" in Eurasia, circa 11.5-11 Ma. Subsequently, the Hipparion .!•!.• and Cremo-
hipparion lineages appear to have undergone major evolutionary radiations in Eurasia
circa 8-6 Ma. The "Sivalhippus Complex" apparently had two separate biogeographic
extensions, the first from Indo-Pakistan to Sub-Saharan Africa (circa 8 Ma), and the
second into China and Europe (circa 5 Ma). Both of these deployments yielded subse-
quent provincial diversification of hipparions. Whereas the Chinese radiation appears
to have been of a cladogenetic sort, the African and European patterns are presently
too poorly characterized to ascertain their evolutionary mode.
The clustering of biogeographic extensions and evolutionary radiations around

the 11 Ma, 8 Ma, and 5 Ma intervals is in itself intriguing. Haq et al. (1988) have
shown, as have earlier authors working with the deep sea-based paleoclimatic record,
that these are intervals of major global climatic change. Vrba (1985) stated that when
good phylogenetic records are accompanied by good geochronologic records, one could
predict globally simultaneous punctuations in evolutionary events and extinctions.
Bernor (1985b) countered that while global climatic events may directly effect
moments of evolutionary radiation, provincial paleogeographic and climatic conditions
may either promote or buffer effects of global events. Despite their limited phylo-
genetic resolution, Old World hipparions would appear to have undergone evolutionary
pulses at distinct intervals of paleoclimatic change. This observation gives credence
to the central core of Vrba's hypothesis: global climatic change, as measured by deep
sea records, may have had a direct and measurable effect on the timing and magnitude
of mammalian community evolution. One could add to this, of course, that such
events directly facilitated biogeographic extensions by exposing previously submerged
land areas, or shifting patterns of provincial climates, and in themselves offered new
opportunities for adaptation by mammalian (here hipparionine horses) lineages. The
patterns cited here certainly provide an intriguing basis for further hypothesis testing.
CONCLUSIONS
We have used 43 of the better known and characterized species of Old World
hipparionines, their North American sister- and outgroup species, to seek refinement
of their evolutionary relationships. Evolutionary analysis of hipparions is always
complicated by morphological variability due both to ontogeny and individual varia-
tion, frequent diversity of species in a single locality, and problems stemming from
the use of discrete or continuous data. However, we provide evidence here that
Eurasian and African hipparionines include several distinct lineages of likely generic
rank. Whether all hipparionines are best included in a single tribe remains to be
determined.
Our analysis here has focused on the use of discrete data. However, we advo-
cate the use of continuous data also, but within a carefully organized research
program whereby a single quarry sample of complete articulated skeletons, belonging
to a single species, is thoroughly analyzed statistically as a population sample. This
work is currently being realized with the study of the H~wenegg hipparion sample
from Hegau, West Germany. When this study is completed, investigators should be
able to more certainly determine the number of species in a sample using both contin-
uous and discrete morphological data.
315
We have attempted to represent current debates concerning the origin of the
various Old World superspecific groups as hypotheses in need of further testing. We
wish to further qualify that the character states used here are in need of further
refinement, and new character states should be sought and incorporated into subse-
quent analyses. Use of continuously distributed data will also undoubtedly prove
useful for future refinements of hipparionine evolutionary history. As a more refined
phylogeny and chronology of these horses is realized, so will our understanding of their
diversity and paleobiology.
ACKNOWLEDGMENTS
We thank Drs. E. Lindsay, V. Fahlbusch, and P. Mein for inviting us to make this
contribution. We thank Dr. Franz Sturmer, Vienna, for his superlative efforts in
preparing the illustrations and Dr. Walter Mittman, Karlsruhe, for his help in calcu-
lating the statistics and preparing the bivariate plots included in figure 4. We would
like to acknowledge financial support to Bernor from NSF (Grant BSR-851736), the
Alexander Von Humboldt Stiftung for his study in Germany during the preparation of
this paper, and Howard University College of Medicine for his sabbatical support;
support to Bernor and Tobien from NATO (Grant No. RG 85/0045) for study of central
European hipparions; support to Bernor, L.-E. Hayek, and Woodburne (NSF Grant No.
BSR 88-06645) for study of North American and Old World hipparion phylogenetic
relationships. We also wish to thank Drs. Richard Hulbert and Bruce MacFadden for
providing us with yet unpublished information on merychippine evolution so critical for
our evaluation of hipparionine evolutionary relationships.
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319
THE HIPPARIONS OF THE LOWER AXIOS VALLEY
(MACEDONIA, GREECE). IMPUCATIONS FOR THE NEOGENE
STRATIGRAPHY AND THE EVOLUTION OF HIPPARIONS
George D. Koufos
Department of Geology and Physical Geography

Aristotelian University of Thessaloniki
P.O. 351-1
54006 Thessaloniki, Greece
INTRODUCTION
The Neogene mammalian localities of the lower Axios valley were discovered at
the beginning of this century and there are several references about them (Andrews,
1918; Bourcart, 1919; Arambourg and Piveteau, 19Z9). The best of the old collections,
now stored in Paris (Museum National d'Histoire Naturelle), is that of Arambourg, who
collected the material during his army service in Macedonia during the First World
War. After that collection nobody excavated in the area and only some isolated
specimens were found by the field workers. In 197Z the University of Thessaloniki
(Laboratory of Geology and Paleontology) in collaboration with the University of Paris
VI and later with the University of Poitiers (Laboratoire de Paleontologie des
VertE~bres et Paleontologie Humaine) began a new campaign of excavations in the
lower Axios valley. Several mammalian localities were discovered and abundant
material has been unearthed. Most of the excavated sites are new, as those of RPl,
RZ/1, PXM, VTK, DIT, DKO. The new localities, the new collected material, its
determination and comparison with those of Eurasian localities allow a detail dating of
the Neogene deposits of the lower Axios valley. The research and the excavations in
the area have continued and always new and interesting material is unearthed.
All the known localities of the lower Axios valley are situated on both sides of
the river Axios and the most important of them are given in figure 1. Most of them
are located in the eastern bank of Axios around the villages of Nea Mesimvria,
Vathylakkos, Prochoma, and Pentalophos. Three localities are known from the
western bank of Axios and they are situated between the villages of Dytiko and Athyra
(figure 1). The localities RZO, VAT, and DTK are referred by Arambourg and
Piveteau (19Z9) under the names "Ravin des Zouaves," "Ravin de Vatili.ik," and "Ravin
de Konikovo," respectively.
The collected material from these localities contains abundant hipparions, which
have been studied during the last ten years. The first collection of hipparions with
few specimens was studied by Koufos (1980). A more detailed study of this materi<~.l,
as well as of that collected after 1980, was begun in 1985 (Koufos, 1985, 1986,
1987a,b, 1988, in press). The results of this study, the dating of the localities cmd of
the Neogene formations with hipparions, the Hipparion biozones in the area, and the
evolution of the lower Axios valley hipparions are given in the present article.

Edited by E. H. Lindsay et al.
Cot)
....,
....,
Xirochorion
GIANNITSA
•
Koufalia e
j~._.~-·-· ---·,.-......
-.- .....__,..._;
0 2 4 6km
I I I I
Fig. 1. Sketch map with the Neogene mammalian localities of the lower Axios valley. RPl = •Ravin de la
Pluie,• RZ/1 = •Ravin des Zouaves-1,• RZO = •Ravin des Zouaves-5,• PXM = •Prochoma-1,• VLO =
•vathylakkos-1,• VTK = •vathylakkos-z,• VAT= •vathylakkos-3,• DTK = •nytiko-1,• DIT = •nytiko-z,•
DKO = •nytiko-3. •
LITHOSTRATIGRAPHY
The lower Axios valley belongs to the Axios (Vardar) zone (Mercier, 1968). The
depressions of the basin were mainly filled by continental deposits. A general paleo-
geographic evolution of the basin is given by Christodoulou (1965). Th~ origin of the
basin is due to a depression of the area during late Eocene. The sea occupied this first
basin until late Oligocene-early Miocene. During that period vertical upli'fting move-
ments caused the regression of the sea, while in Burdigalian the sea entered again in
the basin. During Tortonian the sea regressed again and a large transgression took
place in Sarmatian. After that the sea regressed rapidly and at the beginning of
Pontian the area subsided slowly. Its surface was always above sea level and thus the
erosion was strong and abundant fluvioterrestrial sediments were deposited. These
deposits contain the studied mammalian localities. During early Pliocene the area was
covered by lakes with lacustrine sediments, while in the Pleistocene the southern part
of the area was covered again by the sea. The late Miocene deposits of the lower
Axios valley are divided into two groups; the lower group consisted of red-beds,
gravels, and sands with hipparion-fauna of Pontian age, and the upper group consisted
of white marls and limestones with pollen of Miocene-Pliocene age (Mercier, 1968).
Recently the late Miocene deposits of the lower Axios valley were divided and dated
more precisely. Three different formations were distinguished, described, and dated
with mammalian fauna (Bonis et al., 1985, 1987).
Nea Mesimvria Formation
This formation crops out in the area between the villages of Agionerion, Nea
Philadelphia, Pentalophos, and Nea Mesimvria. It consists of red-beds with abundant
sands and gravels. The red-beds are well indurated and contain a lot of fossils, which
are concentrated into pockets. The fossils were not carried far by the water because
bones in natural connection have been found. Two localities were found in this forma-
tion, "Ravin de la Pluie" (RPl) and "Ravin des Zouaves-1" (RZ/1). Both are situated
north of the village of N. Mesimvria (figure 1).
Vathylakkos Formation
The formation crops out in the area between the villages of Nea Mesimvria,
Prochoma, and Vathylakkos (figure 1). It consists of yellowish marls in the base, which
are overlain by alternating beds of white or yellowish marl, sandy-marl, sand, and
gravels, sometimes with cross-bedding. The locality "Ravin des Zouaves-5" (RZO) is
situated in the yellowish marl of the base while four other localities, Vathylakkos-
1,2.,3 (VLO, VTK, VAT) and Prochoma-1 (PXM), are situated in the above layers,
usually in the sandy marls (figures 1 and Z). There are also some other localities but
with very few fossils. The bones were deposited not very far from the point of their
death because a lot of them have been found in association.
Dytilto Formation
Dytiko Formation crops out in the area of the villages Dytiko and Agrosykea
(figure 1). It consists of alternating beds of white-yellow or gray sand, gravels, sandy
marl, and marl. The top of the formation consists of fresh-water, tuffaceous-massive
limestone, known from the area of the villages Agrosykea and Dassero. Three locali-
ties, Dytiko-1,Z,3 (DTK, DIT, DKO), are known, which are situated along the
Platanoremma River, south to the village of Dytiko (figures 1 and Z). The fossils were
not carried far by the water because bones in natural association were found (e.g.,
hipparion's foot).
Among the known localities of the lower Axios valley there is a site named
"Diavata." A large hyaena, H. salonicae, and some hipparion bones are known from
this locality (Andrews, 1918). Our efforts to find it were unsuccessful but it is prob-
ably situated in one of the ravines between the villages of Diavata, Pentalophos, and
Oreokastron (Koufos, 1985). Recently we found a new locality near the village of
Pentalophos but the available data for its dating, as well as for its correlation with
that of Andrews (1918), are few.
323
LOCALITIES AND FAUNA
The collected material from the above mentioned localities is abundant and
allow us a good identification. The data for the following faunal lists of each locality
were taken from Bonis et al. (1979, 1985, 1986, 1987), Bonis and Koufos (1981, in
press), Bouvrain (1975, 1978, 1980, 198Z, 1986), Geraads (1978, 1979), and Koufos
(1979, 1980, 198Z, 1984, 1985, 1986, 1987a,b,c, 1988, in press).
Ravin de Ia Pluie (RPI)
Progonomys cathalai Samotragus praecursor

Adcrocuta eximia leptoryncha Prostrepsiceros ~.) vallesiensis
Plioviverrops orbignyi Decennatherium pachecoi
Choerolophodon pentelici Palaeotragus coelophrys
Hipparion primigenium Palaeotragus rouenii
Hipparion macedonicum Bohlinia cf. attica
Rhinocerotid uniden. Ouranopithe~acedoniensis
Mesembriacerus melentisi
Ravin des Zouaves-1 (RZ/1)
Ictitherium cf. hipparionum Hipparion sp. (small size)

Adcrocuta eximia Mesembriacerus melentisi
Choerolophodon sp. Ouzocerus gracilis
Hipparion sp. (large size) Samotragus praecursor
Ravin des Zouaves-5 (RZO)
Adcrocuta eximia Prostrepsiceros (Helicotragus) zitteli

Chasmaporthetes sp. Prostrepsiceros (Helicotragus) rotundiconnis
Ictitherium robustum Nisidorcas planicornis
Choerolophodon pentelici Tragoportax rugosifrons
Zygolophodon tapiroides Gazella sp.
Hipparion proboscideum Helladotherium duvernoyi
Hipparion dietrichi Microstonyx maior
Hipparion macedonicum Potamochoerus Postpotamochoerus) hyotheriodes
Dicerorhinus orientalis Mesopithecus n. sp.
Palaeoreas zouavei
Vathylakkos-1 (VLO)
Hipparion dietrichi Nisidorcas planicornis

Palaeoreas lindermayeri
Vathylakkos-Z (VTK)
Ictitherium robustum Tragoportax rugosifrons

Ictitherium hipparionum Gazella sp.
Plioviverrops orbignyi Nisidorcas planicornis
Choerolophodon pentelici Bohlinia attica
Hipparion dietrichi Microstonyx major
Hipparion macedonicum
Vathylakk.os-3 (VAT)
Plesiogulo crassa Samotherium boissieri

Ictitherium robustum ?Helladotherium duvernoyi
Ictitherium hipparionum Bohlinia attica
Plioviverrops orbignyi Gazella sp.
Hipparion dietrichi Tragoportax rugosifrons
Hipparion macedonicum Nisidorcas planicornis
324
Dicerorhinus orientalis Prostrepsiceros (Helicotragus) zitteli
Dorcatherium puyhauberti Microstonyx major
Prochoma-1 (PXM)
Adcrocuta sp. Hipparion macedonicum

Ictitherium sp. Tragoportax rugosifrons
Choerolophodon cf. pentelici Nisidorcas planicornis
Hipparion dietrichi Gazella sp.
Dytiko-1 (DTK}
Chasmaporthetes bonisi Protragelaphus theodori

Choerolophodon pentelici Palaeoreas lindermayeri
Dicerorhinus orientalis "Graecoryx cf. valenciennesi"
Hipparion mediterraneum Gazella sp.
Hipparion matthewi Bovid uniden. (large size)
Hipparion periafricanum Pliocervus pentelici
Bohlinia attica Mesopithecus pentelicus
Dytiko-Z (DIT)
Hipparion mediterraneum "Graecoryx cf. valenciennesi"

Hipparion matthewi Gazella sp.
Bohlinia attica Bovid uniden. (large size)
Palaeotragus rouenii Dorcatherium puyhauberti
Palaeoreas lindermayeri Mesopithecus pentelicus
Dytiko-3 (DKO)
Protictitherium crassum "Graecoryx cf. valenciennesi"

Hipparion mediterraneum Gazella sp.
Hipparion matthewi Mesopithecus pentelicus
Hipparion cf. periafricanum Hystrix sp.
Protragelaphus theodori
Diavata
"Hyaena" salonicae Hipparion sp. (large-medium size)
THE HIPPARIONS OF THE LOWER AXIOS VALLEY
The genus Hipparion was referred in the lower Axios valley by Andrews (1918),
who described a maxilla and some bones under the name "H. gracile." The material
was found near the village of Diavata, about 4 km northwest of Thessaloniki and it is
stored in the British Museum (Natural History). Recently this material was studied,
described, and compared with those of the other Axios localities (Koufos, 1985). It is
a medium to large-sized hipparion with rich enamel plication, large subtriangular
isolated protocone, multiple pli caballin, elliptical hypocone, and well defined lingual
hypoconal groove. These characteristics are primitive and give the idea for a
Vallesian/Turolian form; no specific name was given to this hipparion because of the
few specimens and the inexact location of the material. One year later, Bourcart
(1919) studied the Neogene deposits of the area and he referred to the presence of "H.
gracile" from the area of the village of Vathylakkos. The material collected by
Arambourg was studied later; all the hipparions were referred to the species "H.
gracile" and were considered similar to the Pikermi ones (Arambourg and Piveteau,
1929). Much later, the hipparions of the Arambourg collection were studied by Pirlot
(1956), who noted that their dentition is similar to that of the Pikermi material and
thus he suggested the presence of the species "H. gracile" and H. mediterraneum in
the Axios valley. More later, Forsten (1968) studied the Arambourg collection and she
325
w
....,
(l) Table 1. The lower Axios valley hipparions with their characterist ics.
H. pJtObo~c..idwm H. cUwucJU H. medli:eiVUlnwm H• matthwe..i H.p~

H• p1Wn<.genl.um H. mac.edoniewn
RPl,RZ 1 ,RZO,PXM, RZO.PXM,VLO,
RZO DTK,DIT,DKO DTK,DIT ,DKO DTK, ?DKO
RPl VTK,VAT V.TK,VAT
Small Large Medium Medium Small Very small

SIZE Large
Relatively long Long- Wide Short- Wide Short - Narrow Shol't - Narrow
MUZZLE - and wide.
-
Deep, situated Short, situated
not far in front V~ry deep. Moderately deep Moderately deep far in front of -
NARIAL OPENING -
of the orbits. the orbits.
PREORBITAL Single - Oval Single - Oval Single - Oval - -

- Double
FOSSA Well defined Faint
DISTANCE ORBIT - Short Short Long Moderate - -

PREORB. FOSSA
Rich Rich - Moderate Rich Moderate - Simple Moderate -Simple Simple
ENAMEL PLICAT.
p 3 •4 = 12.4 p 3 • 4 = 14.5 p3,4= 9.0 Very simple
P 3 • 4 = 18.4 p 3 • 4= 12.3 P 3 •4= 17.6
PLICAT. NUMBER
M1 • 2= 17.4 M1 • 2= 12.6 M1 •2
= 16.3 M1 • 2= 10.3 M1 • 2= 12.7 M1 • 2 = 9.0
Elliptical. Rounded- Elliptical. Rounded- Elliptical
Elliptical-Small Blipticsl Elliptical Isolated except Isolated except in Rounded Isolated except
Isolated I
PROTOCONE Isolated Isolated Isolated in the very the very worn P2.
worn teeth. teeth. I
I
DISTAL HYPOCO- Deep Shallow

Deep Deep Shallow Shallow Deep
NAL GROOVE
LINGUAL HYPO- Slightly curved Very slightly Absent Absent Absent Absent Absent
CONAL GROOVE enamel. defined.
---- '-----·-···--·- -
Small - Simple.
Absent or rudime- Small - Simple Rudimentary in P.
Large Small Large - Multiple Small - Simple
PLI CABALLIN ntary in the very Double in P 2 . Absent in M.
Simple - Double Simple -Double (1-4 plisl
worn teeth.
-
Well Well Well Moderately Well Moderately
PARASTYLID -
developed developed developed developed developed developed
Small Very Small Moderate -

PROTOSTYLID - Small
only in M1 . small Rarely present only in M1 •
ENAMEL OF Plicated or Slightly plicated Plicated or Simple Simple Simple -

FLEXIDS Cl'j!nelated. or crenelated. crenelated.
U-shaped V-shaped Shallow in P V-shaped U-shaped U-shaped

LINGUAFLEXID -·
Shallow Shallow Deep in M Shallow Shallow Shallow
\\fell developed Small only in Well developed

PLI CABALLINID in the little the little in the little Absent Usually absent Absent -
worn teeth. worn teeth. worn P.
Brachyodont Hypsodont Hypsodont

p 3 • 4 = 49.8 p 3 • 4 = 39.6 p3,4 = -
M1,2 = M1 • 2 = 41.0 M1 • 2 = 42.7 -
HYPSODONTY - - - -
P3,4 = - P3,4 = - P 3 , 4 = 42.8
M1 , 2 = 46.0 M1 , 2 =41.8 M1 , 2 = 39.2
METAPODIALS Short-Robust Long-Slender Robust Long- Slender Long-Slender Long-Slender -
w
N
.....
considered that two hipparions are present in Axios valley, the medium-sized H.
mediterraneum and the small-sized H. matthewi. Recently the skulls of hipparions in
the Arambourg collection were studied by Woodburne and Bernor (1980) and, according
to the morphology of the preorbital fossa, they were assigned to Group Z and 3, while
the presence of Group 4 is possible. One year later, Woodburne et al. (1981, figure Z)
identified the species H. mediterraneum, H. matthewi, and H. cf. prostylum from the
Arambourg collection.
I have seen all the hipparions of the Arambourg collection in the Museum
National d'Histoire Naturelle of Paris. Unfortunately, the material is mixed and there
are no locality indications on the fossils. Thus, it is difficult to distinguish the differ-
ent localities (six different localities with Hipparion were mentioned by Arambourg
and Piveteau, 1929) to which the material belongs. The type of fossilization and the
color of the fossils indicate that most of the material is from Vathylakkos-3 (VAT). I
have not worked enough on this material because I wanted first to finish with the new
collected material and then to use this for comparison with the older material.
Nevertheless, I think that in the old material there are at least three species: a
medium-sized (possibly H. dietrichi), a small-sized (probably H. macedonicum), and a
large-sized (there are two mandibles of a large hipparion). It is also possible that the
presence of H. matthewi is represented in the material from Dytiko but the mixing of
the material makes the distinction difficult. I hope that the detailed study and
comparison of this material with the new and stratigraphically controlled material will
give satisfactory data about its provenance and species number.
The unearthed material during our excavations in the lower Axios valley contains
abundant hipparions, found in various localities and stratigraphic levels. This well
located Hipparion material allows us to determine and compare this, to find the rela-
tions between the determined species, to date the deposits, and to divide them in
biozones based on Hipparion. The determined Hipparion species of the lower Axios
valley, the localities in which they were found, the stratigraphic horizons they are
assigned to, and a small description with the main characteristics of each species are
given in the following (see also table 1).
Bipparion primigenium (von Meyer, 18Z9)
Locality: RPl
Horizon: Late Vallesian (late Miocene), MN-10
Description: The studied material contains a mandible, some mandibular pieces,

isolated teeth, and some bones described by Koufos (1986). It is a large-sized
hipparion with richly plicated upper cheek teeth, elliptical and isolated protocone,
well developed hypocone with deep distal hypoconal groove, and slightly curved
enamel in the position of the lingual hypoconal groove. The lower cheek teeth have
well developed parastylid, plicated, and crenulated enamel in the flexids border, well
developed pli caballinid in the little worn teeth and shallow U-shaped linguaflexid.
The teeth seem to be brachyodont (high hypsodonty index, table 1). The metapodials
are short and robust, but more slender than those of Eppelsheim. The comparison of
H. primigenium from Axios valley with other Vallesian hipparions from Eurasia shows
that it is more advanced than the early Vallesian forms (Eppelsheim, Nombrevilla, Can
Llobateres) indicating a late Vallesian age for it (Koufos, 1986).
Bipparion macedonicum Koufos, 1984

Locality: RPl, ?RZ/1, RZO, VTK, VAT, PXM
Horizon: Late Vallesian-early Turolian (late Miocene), MN-10, 11
Description: The species was first described from RPl (Koufos, 1984, 1986) and
later it was found in the other referred localities (Koufos, 1987a,b, 1988). It is a
small-sized hipparion with a relatively long and wide muzzle and elongated narial
328
opening situated not far in front of the orbits. The preorbital fossa is single, oval,
well defined, and it is situated near the orbits. The upper cheek teeth have rich-
moderate enamel plication, elliptical and isolated protocone, simple-double pli
caballin, well developed hypocone with well developed distal hypoconal groove and
slightly defined lingual hypoconal groove. The lower cheek teeth have a well
developed parastylid, small protostylid, small pli caballinid in the little worn
premolars (in the molars it is absent), slightly plicated and crenulated enamel in the
flexids and shallow V-shaped linguaflexid. The available and measurable teeth
indicate hypsodont teeth (table 1). The metapodials are elongated and slender. The
comparisons with the other known small-sized hipparions show that H. macedonicum is
distinct and represents the oldest known small-sized hipparion in Europe.
Hipparion proboscideum Studer, 1911
Locality: RZO, ?PXM, ?VTK
Horizon: Early Turolian (late Miocene), MN-11
Description: It is known from a frontal part of the skull, a piece of mandible,

some isolated teeth and some post-cranial bones, all from RZO (Koufos, 1987a). There
are also some isolated bones of a large-sized hipparion from PXM and VTK, which
possibly belongs to H. proboscideum (Koufos, 1987b, 1988). It is a large-sized
hipparion with long and wide muzzle, very deep narial opening, and double preorbital
fossa situated near the orbits. Both fossae are elliptical, the lacrymal is deep and well
defined; the nasal fossa is shallower and its anterior part is not well defined. The
upper cheek teeth have rich enamel plication, elliptical and isolated protocone,
multiple pli caballin and elliptical hypocone with only a shallow distal hypoconal
groove. The lower cheek teeth have well developed parastylid, small protostylid only
in M/1, slightly plicated or crenulated enamel in the flexids, well developed pli
caballinid in the premolars and rudimentary in the molars, and a linguaflexid which is
shallow in the premolars and deep in the molars. The metapodials are robust.
Hipparion dietrichi (Wehrli, 1941)
Locality: RZO, VLO, VTK, VAT, PXM
Horizon: Early Turolian (late Miocene), MN-11
Description: There is a lot of material of this species from the various locali-
ties and allow us a good description (Koufos, 1980, 1987a,b, 1988). It is a medium-
sized hipparion with short and wide muzzle, moderately deep narial opening with
single, faint, and elliptical preorbital fossa, which is situated far from the orbits. The
upper cheek teeth have moderate-simple enamel plication, elliptical and isolated
protocone, which in the very worn teeth is connected with the protoloph, small and
simple pli caballin, and elliptical hypocone with shallow distal hypoconal groove only.
The lower cheek teeth have a moderate parastylid, very small protostylid, no pli
caballinid, simple enamel in the flexids, and V-shaped shallow linguaflexid. The meta-
podials are long and slender. The calculated hypsodonty index for the upper and lower
cheek teeth (table 1) indicates that the teeth are hypsodont. The comparisons with
the material from other localities indicate that it is similar to the material of H.
dietrichi from Samos, Q/1-4.
Hipparion mediterraneum (Roth & Wagner, 1855)
Locality: DTK, DIT, DKO
Horizon: Late Turolian (late Miocene), MN-13
Description: There is enough material from the Dytiko localities which helps us
to identify the morphological characters of this species. It is a medium-sized
hipparion with short and narrow muzzle, moderate narial opening, single oval pre-
329
orbital fossa, and moderate distance orbit-preorbital fossa. The upper cheek teeth
have moderate-simple enamel plication, a rounded-elliptical and isolated protocone,
which in the very worn teeth is connected with the protoloph, small and simple pli
caballin which is rudimentary or absent in the very worn teeth. The lower cheek teeth
have a well developed parastylid, small and rarely present protostylid, simple enamel
in the flexids, usually no pli caballinid, and U-shaped, shallow linguaflexid. The
metapodials are long and slender. It is very similar to the Pikermi H. mediterraneum
(Koufos, in press). -
Hipparion matthewi Abel, 19Z6
Locality: DTK, DIT, DKO
Description: This is a small-sized hipparion with short and narrow muzzle and
short narial opening, which is situated far in front of the orbits. The upper cheek
teeth have simple enamel plication, a rounded and isolated protocone, small, simple pli
caballin (double in PZ/), and elliptical hypocone with well developed distal hypoconal
groove. The lower cheek teeth have a moderately developed parastylid, moderate
protostylid in M/1 only, simple enamel in the flexids, no pli caballinid, and shallow,
U-shaped linguaflexid. The metapodials are elongated and slender.
Hipparion periafricanum Villalta &: Crusafont, 1957
Locality: DTK, ?DKO
Description: The available material is not abundant; two pieces of maxilla and
some bones, which possibly belongs to this species (Koufos, in press). It is very small
sized and the upper cheek teeth have closed and isolated fossettes (except PZ/ in
which they are connected). The enamel plication is very simple, the boucle-
prefossette (Sondaar, 1961) is absent and the pli caballin is rudimentary in the pre-
molars and absent in the molars. The protocone is rounded-elliptical and isolated,
except in PZ/. The pli protoloph and pli hypostyle (Sondaar, 1961) are usually absent,
especially in the molars. The metapodials are elongated and slender. The comparisons
with the type material from Spain show that the teeth are morphologically and
metrically similar with the Dytiko material. The bones from Dytiko are larger than
those from the type locality and thus they are referred to as H. cf, periafricanum
(Koufos, in press).
BIOSTRATIGRAPHY
The identification and the comparisons of the lower Axios valley hipparions
provide data for dating of the deposits and for their division into biozones. Moreover,
the morphological characters of the hipparions can help us with the dating and how
these horses changed during geological time. The rest of the fauna from each locality
can also help to test the Hipparion dating.
Nea Mesimvria Formation is the oldest stratigraphically and contains two

species, H. primigenium and H. macedonicum. The first species indicates Vallesian
age. The type locality for H. primigenium is Eppelsheim (Germany) which is of early
Vallesian age. The RPl material, with less plicated and more hypsodont cheek teeth,
longer and more slender metapodials, seems to be younger than Eppelsheim (Koufos,
1986). Moreover, in the early Vallesian localities (Eppelsheim, Nombrevilla, Can
Llobateres) only one Hipparion species is known, while in the younger localities of late
Vallesian (Masia del Barbo, Montredon) more than one species are known (Sondaar,
1974; Eisenmann, in press). Thus, RPllocality with two hipparions seems younger than
early Vallesian. The rest of the fauna contains some characteristic species, which
330
Table z. Plication number of the lower Axios valley hipparions (all the
available hipparion's teeth are counted together and the mean
plication number for each locality is given).
LOCALITIES HIPPARION PLICATION NUMBER

SPECIES Premolars Molars
DKO 3 11.0 9.4

DIT 2 13.3 13.0
DTK 3 13.0 11.7
VAT 2 12.5 9.8
VTK 3? 12.7 10.6
PXM 3? 12.4 11.6
RZO 3 11.9 11.9
RPl 2 17.1 15.9
confirm the late Vallesian age of the Nea Mesimvria Formation. The hyaena A.
eximia leptoryncha is more primitive than the typical A. eximia from Pikermi, indi-
cating an older age (Bonis and Koufos, 1981). Mesembriacerus is a primitive ovibovine
differing from Turolian ovibovines from China and Samos (Bonis et al., 1985, 1987).
The rodent Progonomys cathalai, which is absent in Nombrevilla and perhaps in Can
Llobateres, is frequent in Masia del Barbo. Its presence in RPl indicates a late
Vallesian age. Thus Nea Mesimvria Formation is older than Turolian and a later
Vallesian age is possible for it; more precisely its age may be more than 9 Ma, while
RPl and RZ/1 may be between 9-11 Ma (Bonis et al., 1987).
The hipparions of the Vathylakkos and Dytiko Formation have more advanced
characters than those of the Nea Mesimvria Formation and seem to be younger. The
enamel plication [by the plication number of Forsten (1968)] is higher in the Vallesian
than in the Turolian hipparions except for the Turkish forms (Sen et al., 1978; Staesche
and Sondaar, 1979). The Vathylakkos and Dytiko hipparions have less plicated upper
teeth than those of Nea Mesimvria (table 3) and in that feature they seem to be
younger and more closely related with the Turolian forms. The hypsodonty of
hipparion's cheek teeth can also give data about their relative age. The hypsodonty is
given by the hypsodonty index, which is calculated by dividing the length of the tooth
(at the middle of its height) by the height of the tooth. The cheek teeth of hipparions
are more brachyodont in the primitive Vallesian forms and more hypsodont in the
Turolian ones (Sondaar, 1961; Forsten, 1968; Sen et al., 1978). The calculated hyp-
sodonty indices for the hipparions of Axios valley are given in table 1; measurable
teeth were few and thus these values can give only suggestions about the hypsodonty.
However, the teeth of Vathylakkos and Dytiko hipparions appear more hypsodont than
those of Nea Mesimvria, indicating younger age.
Sen ;t al. (1978) considered the keel development as a morphologic character

distinguiShing the Vallesian from the Turolian hipparions. They used a keel index (DAP
keel x 100/DAP min. art. distal) and found that this index increases from Vallesian to
Turolian. The keel index of the Axios valley hipparions (table 3) clearly increases in
the metatarsals from the Nea Mesimvria to Vathylakkos and Dytiko hipparions;
although the differences in the mean values are not very high. In the metacarpals of
the lower Axios valley hipparions one can say the opposite, that the keel index
decreases from the Nea Mesimvria to Vathylakkos and Dytiko hipparions. However,
the small number of metacarpals may not give definitive results or the keel develop-
ment may be a variable feature and cannot be used as evidence for the age of
hipparions. The keel index of the metatarsals indicates that the Vathylakkos and
Dytiko hipparions are younger than those of the Nea Mesimvria.
331
Table 3. Keel index for the metapodials of the lower Axios valley
hipparions.
LOCALITIES n Melli n Mtlll
RPI 2 120.0 2 120.0

RZ 1 - - 1 122.2
RZO 1 118.9 8 126.6
PXM 11 119.6 10 123.7
VTK - - 1 125.6
VAT 5 122.3 4 125.1
DTK 13 119.9 28 126.4
DIT - - 3 130.3
DKO 4 116.6 3 127.7
Moreover, the Hipparion species and the rest of the fauna help us to correlate
the Vathylakkos and Dytiko Formation. The presence of H. macedonicum in
Vathylakkos Formation indicates a late Vallesian age. The other two species, H.
proboscideum and H. dietrichi belong to the third Hipparion zone of Mediterranean
Neogene, indicating early Turolian (Sen et al., 1978). Both species are known from
Samos Q/1-4, which are considered older than Pikermi. Thus, the Vathylakkos
hipparions indicate a latest Vallesian-early Turolian age. The rest of the fauna of the
formation supports this age. The presence of Dorcatherium, the antelope Nisidorcas
planicornis known from Vallesian/Turolian localities (Kayadibi) and the smaller size of
Prostrepsiceros (H.) rotundicornis relative to specimens of Pikermi indicate that the
Vathylakkos fauna is older than Pikermi, and more similar to Samos Q/1-4 (Bonis et
al., 1987). Thus the Vathylakkos Formation age is considered latest Vallesian-early
Turolian, and can be estimated between 7-9 Ma. The hipparions of Dytiko Formation
include two small-sized species (H. matthewi, H. periafricanum), which are character-
istics of the fourth Hipparion zone of Mediterranean Neogene, indicating late Turolian
age (Sen et al., 1978). Thus, the Dytiko hipparions indicate a late Turolian age. The
rest of the fauna and especially Protragelaphus and Miotragocerus show characters
clearly more advanced than those of Pikermi, supporting the idea for a younger age
(Bonis et al., 1985, 1987). Palynological study of the limestones, situated at the top of
the formation, gave elements of Miocene/Pliocene age (Mercier and Sauvage, 1963).
All the available data for the Dytiko Formation indicate a late Turolian age for it.
Considering the Miocene/Pliocene boundary as being 5.5 Ma, then the Dytiko
Formation age is placed about 7-5.5 Ma.
The exact stratigraphic position of the Diavata locality was questionable

because of the few known specimens and the unknown site in which it was found.
Recently the study of the available material and its comparison with other well known
samples gave enough data for the age of the locality. The morphological characters
and the metrical comparisons of Hipparion remains indicate a form similar to the
Vallesian/Turolian ones (Koufos, 1985). The giant hyaena ("Hyaena" salonicae) from
the Diavata locality seems similar to the large percrocutoid hyaenas and has a
morphologically primitive appearance (Howell and Petter, 1985). During the last year
we found a new locality Pentalophos-1 (PNT) near the village of Pentalophos, in which
we found the mandible of a giant hyaena. Unfortunately, the comparison with H.
salonicae is impossible because this species is known only from a maxilla. However,
the neighboring areas from which both specimens have come and their size allow us to
suppose some similarities between them. Thus, a late Vallesian or early Turolian age
for the Diavata locality is possible.
332
CONTINENTAL "' HlPPARlON
Ma 3w LI THOSTRATIGRAPHY L OCALI TIES f A U N A
SERIES/STAGES ~ ~ 810ZONES
H..Q:IJ)41U.on r~e.d.U.eJVt.ane.um
% H.ippalt..ion ma.UI.vei
0
i
H4>pall.iou pw%JUcauum
Clut6mapo.u.Jte..tu bon..i.6.i
z 13 - UOilca.thWUn! w!{ftaub<JLU
P~totAagelapluLb .t.he.odclti.
Bol.t.itU:a att..:ca
w Oyt1ko-l, 2 , 3
z
Ul 12
.J
0
0
..
0
H<ppall.iou cU.e.tMcJU:
H.ip(J(llt.ion macedott..i.c.um
.... 2
0
H<ppa!Uou pwbo~cUI•um
11
i ~ Va thylakkos-1,2 ,3
1et<..thwum ~o~~um
1c.ti.tlte.lli.wrt IUppa.ll...icJtum
Prochoma-1 Vo"-"'Lti•wum puflhaub<JLU
N.U..idCJtCM p£att.-i.cOJtlt.i4
Ul 1- Pa.iae.CJte£1.6 zouavei
hagopolt.ttu wgo~~o""
1- Rav in des Zouaves- 5 P~o~~~~ceJt.o~ IH.·I z~eU
H..ippa-Uon pWntge.n.iwn
Hip.oaM.on mo.cWou.icum
Ravin des Zcuaves.-1
10 .J z PMgonomy~ ca.tha.fa.<
...<l:
(/)
10
Ravin de la Pluie Ad~o~ vWnia l'opto"!(ucl•a
Mu<mb.UoCeA~ mel£~~
Ul
.J OUZOCe/:.0.6_gJLac..il..i.o
.J Oece•ma.tlte.tiwn padtecoi
<l:
>
Fig. z. Stratigraphy of the late Miocene deposits of the lower Axios valley
(Macedonia, Greece).
The various Hipparion species in the different stratigraphic levels of the lower
Axios valley and their morphological characters help us to distinguish some Hipparion
biozones in the late Miocene deposits of the valley. At least two Hipparion species
are known from each locality, a small-sized and a large- or medium-sized species.
Thus, two Hipparion species will be used for the designation of each proposed biozone.
Biozone of B. primigenium - B. macedonicum
Type locality: "Ravin de la Pluie" (RPl), lower Axios valley, Macedonia, Greece
Referred localities: "Ravin des Zouaves-1" (RZ/1), lower Axios valley
Horizon: Late Vallesian (late Miocene)
Diagnosis: Short and wide muzzle, deep narial opening, rich enamel plication in
the upper cheek teeth, well developed lingual hypoconal groove or slightly curved
enamel in its position. The lower cheek teeth have plicated or crenulated enamel in
the flexids and usually the pli caballinid is present. The teeth are brachyodont (high
333
hypsodonty index) and the index (length M/1-M/3 x 100/length P/Z-P/4) is higher than
100; in the RPl material "mean is 105 for H. macedonicum and 101.5 for H.
primigenium (Koufos, 1984, 1986). The metapodials of the large-sized form are short
and robust while in the small-sized species they are long and slender.
Characteristic fauna: Progonomys cathalai, Adcrocuta eximia leptoryncha,

Mesembriacerus melentisi, Ouzocerus gracilis, Decennatherium pachecoi,
Ouranopithecus macedoniensis
Biozone of B. dietrichi - H. macedonicum
Type locality: Prochoma-1 (PXM), lower Axios valley, Macedonia, Greece
Referred localities: "Ravin des Zouaves-5" (RZO), Vathylakkos-1,Z,3 (VLO,

VTK, VAT), lower Axios valley
Horizon: Latest Vallesian-early Turolian (late Miocene)
Diagnosis: Short and wide muzzle, moderately deep narial opening, single,
faint, and elliptical preorbital fossa. The upper cheek teeth have moderate enamel
plication, simple and small pli caballin, and lack the lingual hypoconal groove. The
lower cheek teeth have simple enamel in the flexids, V-shaped linguaflexid, and
usually lack a pli caballinid. The teeth are hypsodont (small hypsodonty index, table 1)
and the metapodials are long and slender. A third large-sized species H. proboscideum
is also present in this biozone.
Characteristic fauna: Plesiogulo crassa, Ictitherium robustum, Ictitherium

hipparionum, Dorcatherium puyhauberti, Nisidorcas planicomis, Palaeoreas zouavei,
Plioviverrops orbignyi, Tragoportax rugosifrons, Prostrepsiceros (Helicotragus) zitteli
Biozone of B. mediterraneum - H. matthewi
Type locality: Dytiko-1 (DTK), lower Axios valley, Macedonia, Greece
Referred localities: Dytiko-Z,3 (DIT, DKO), lower Axios valley
Horizon: Late Turolian (late Miocene)
Diagnosis: Short and narrow muzzle, moderately deep narial opening and single
oval preorbital fossa. The upper cheek teeth have simple enamel plication and simple,
small, or rudimentary pli caballin. The lower cheek teeth have simple enamel in the
flexids, U-shaped linguaflexid, and lack a pli caballinid. The teeth are hypsodont
(small hypsodonty index) and the metapodials are long and slender.
Characteristic fauna: Chasmaporthetes bonisi, Dorcatherium puyhauberti,

Protragelaphus theodoti, Bohlinia attica
EVOLUTION OF THE LOWER AXIOS VALLEY IUPPARIONS
The lower Axios valley hipparions with many species from different stratigraphic
levels and well established biozonation provide us data to suggest some evolutionary
lines for them. The early Vallesian hipparions are still unknown in Greece. The oldest
known hipparions are those of "Ravin de la Pluie" in Macedonia. Some teeth and bones
of a large-hipparion, similar to H. primigenium, are known from the Vallesian locality
of Kastellios (de Bruijn et al., 1971). Some isolated teeth of hipparions from
Halmyropotamos (Evia) and Chalkoutsi (near Athens) are found with H. mediterraneum
and according to their dimensions and rich enamel plication they are identified as the
early Vallesian species H. koenigswaldi (Melentis, 1967; Koumantakis, 1971). I have
not seen this material but I think that it probably belongs to the large-sized H.
brachypus of Pikermi, which has richly plicated teeth and large dimensions (Koufos,
334
1987d). The presence of an advanced H. pr1m1genium in RPl and of a large-sized
hipparion similar to H. primigenium in Kastellios (Crete) indicate that H. primigenium
was possibly present in the early Vallesian of Greece. H. primigenium is known in the
countries neighboring Greece. It is referred from the Vallesian l~calities of Bulgaria
(Nesebr, Kalimantsi, Kromidovo, Ivanane, and Hrabrsko) (Forsten, 1978). In Asia
Minor a large-sized hipparion is known from the Vallesian localities of Yassioren and
Esme-Akcakoy, under the name H. ankyranum. The species is referred to the
Vallesian of Europe and Asia under various specific names and it is considered as the
oldest European hipparion group. In late Vallesian the primitive hipparions began to
differentiate and several forms appeared. The late Vallesian localities usually contain
two Hipparion species. Sondaar (1974) identified two hipparions from Masia del Barbo
(Spain). The late Vallesian locality of Montredon (France) contains a medium-sized
hipparion, named H. depereti (Sondaar, 1974). Recently a small-sized hipparion was
found in the Montredon material (Eisenmann, in press). In the Vallesian/Turolian
locality of Kayadibi (Asia Minor) two hipparions, a large-sized and a medium-sized
species are distinguished (Staesche and Sondaar, 1979). Thus, late Vallesian is char-
acterized by the appearance of the various size groups of Hipparion. The large-sized
group has continued to exist while the branches of the medium-sized and the small-
sized hipparions began to evolve. "Ravin de la Pluie" represents an important stage in
the evolution of hipparions because of the appearance of the small-sized group. In the
known Vallesian localities only large- or middle-sized hipparions were known. H.
macedonicum is the first small-sized representative of that period and it is considered
as the earliest occurrence of the small-sized group. The last forms were known only
from the late Turolian of Eurasia with the specific names H. matthewi, H. gromovae,
and H. periafricanum. Now the late Vallesian H. macedonicum indicates that the
branch of the small-sized hipparions appears earlier, at about the same time as the
medium-sized ones. This early appearance of the small-sized hipparions is confirmed
by the presence of a similar hipparion in Montredon (Eisenmann, in press).
The branch of the large-sized hipparions continued to exist in early Turolian of

the Axios valley with the species H. proboscideum, which is present in the Vathylakkos
localities. This large hipparion is known from Samos Q/1-4 (Sondaar, 1971), and it is
possibly present in Maragheh (Campbell, 1980). The large H. brachypus from Pikermi
possibly belongs to this branch of the large-sized hipparions. The small H.
macedonicum continued its presence during early Turolian, representing the small-
sized branch. The medium-sized hipparions are unknown in the late Vallesian of
Macedonia. Nevertheless, there is a piece of metapodial whose dimensions are
between that of H. primigenium and H. macedonicum (Koufos, 1986, figure 5, table
9). This sole specimen, as well as the unsufficient knowledge of the variations in the
dimensions of both RPl species, is a weak argument for the presence of a third
medium-sized species in late Vallesian. In early Turolian the medium-sized branch is
well known by the presence of H. dietrichi, which possibly came from the A.
primigenium group. This species is also present in the same age localities of Samos
Q/1-4 (Sondaar, 1971) and Maragheh (Campbell, 1980). The medium-sized H.
mediterraneum, which is probably present in the late Turolian of Axios valley, possibly
belongs to the same branch, with H. dietrichi. H. mediterraneum is also present in
Pikermi (middle Turolian) while it is absent in the older localities of the Vathylakkos
Formation and Samos (Q/1-4). Thus, it probably appeared at the end of early Turolian
from the medium-sized line of hipparions. There are no representatives of the large-
sized hipparions in the late Turolian of Axios valley. The small-sized hipparions were
represented in late Turolian by H. matthewi. This species is lmown from Q/ 5 of Samos
(Sondaar, 1971), from Asia Minor (Staesche and Sondaar, 1979), and Maragheh
(Campbell, 1980), always from late Turolian localities. It probably arises from the
branch of the small-sized hipparions and possibly from H. macedonicum. Among the
late. Turolian specimens from the Axios valley there is a very small hipparion similar
to H. periafricanum. This was possibly derived from the H. matthewi group. H.
eriafricanum is known from Spain and is considered to arise from the small-sized H.
gromovae Alberdi, 1974). The last species does not much differ from H. matthewi. In
any case, H. periafricanum is derived from a small-sized hipparion, H:-matthewi or H.
gromovae.
335
This interpretation of the hipparions of the lower Axios valley allows us to
consider that:
1. They came from a large-sized primitive hipparion of early Vallesian, unknown in

Greece.
z. In late Vallesian this primitive group was differentiated and three Hipparion groups
appeared. The large-sized group continued, while a medium-sized and a small-
sized group began to develop.
3. H. macedonicum represents the first representative of the small-sized group; it

appeared in late Vallesian.
4. Three groups continued their presence in the younger intervals by various specific
names.
5. Some very small-sized hipparions were derived from the small-sized group, as H.
periafricanum, in late Turolian.
CONCLUSIONS
The lower Axios valley mammalian localities provide much data about the late
Miocene deposits and the hipparions found in these localities allow us to date the
deposits and establish a biozonation, plus provide data about the evolution of
hipparions. Three formations can be distinguished in the late Miocene of the lower
Axios valley.
a. Nea Mesimvria Formation, with H. primigenium and H. macedonicum of late

Vallesian age.
b. Vathylakkos Formation with H. dietrichi, H. proboscideum, and H. macedonicum of

latest Vallesian-early Turolian age~
c. Dytiko Formation with H. mediterraneum, H. matthewi, and H. periafricanum of

late Turolian age. -
The identified hipparions help us to distinguish three Hipparion biozones.
a. Biozone of H. primigenium- H. macedonicum of late Vallesian age.
b. Biozone of H. dietrichi- H. macedonicum of latest Vallesian- early Turolian age.
c. Biozone of H. mediterraneum - H. matthewi of late Turolian age.
In late Vallesian the primitive H. primigenium group was differentiated and the
medium- and small-sized branch of hipparions appeared. H. macedonicum from RPl is
the first representative of the small-sized group. In early Turolian of the Axios valley
three groups (large, medium, small) of Hipparion are present while in late Turolian
there are only representatives of medium- and small-sized groups.
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338
THE GENUS EQUUS IN EUROPE
A. Azzaroli
Museo di Geologia e Paleontologia

Universita di Firenze
Via la Pira 4
Firenze, Italia
THE ORIGIN OF EQUUS
The genus Equus originated in North America and was derived from the late
Hemphillian (late Miocene to earliest Pliocene) Dinohippus (Quinn, 1955). All the
Pliocene, Pleistocene, and living monodactyl equids of North America and of the Old
World - with the only exception of the possibly early Pliocene Astrohippus stocki
Lance 1950 from Mexico- may be united in the genus Equus. Some authors, notably
Quinn (1955, 1957), Nobis (1971), and Samson (1975), divided them into a variety of
genera and subgenera. This practice will not be followed here and the generic name
Eguus will be used in the broad, traditional sense. This is because, in spite of signifi-
cant variation among equids of different ages and geographic ranges, their basic
structure remained remarkably uniform.
The genus Equus was assumed to be diphyletic by Bennett (1980) and by Dalquest
(1987), mainly on evidence from the dentition. Bennett assumed a derivation from two
distinct, but not specified species of Dinohippus; there is, however, no reliable
evidence in the fossil record of two distinct species of Dinohippus in the late
Hemphillian, nor of any species of Equus other than ~· simplicidens in the early
Blancan, and dental features advocated by Bennett for a diphyletic origin are of
limited taxonomic significance (Azzaroli and Voorhies, in prep.). Dalquest assumed a
derivation of zebras and horses s. str. from Dinohippus and of asses from Astrohippus,
but apart from the lack of fossil evidence for the critical period comprised between
the last occurrence of Astrohippus in the latest Hemphillian of Mexico and the first
occurrence of possible, but far from proven, ancestors of hemiones and asses in the
late Blancan of the southern United States (Azzaroli and Voorhies, in prep.), Dalquest's
assumption contrasts with the complicated cranial features of Astrohippus, recently
illustrated by MacFadden {1984).
Figure 1 reproduces a skull of Equus simplicidens Cope, the oldest known species
of Equus. The specimen, LACM/CIT 210/17047, comes from the early Blancan
Hagerman fauna of Idaho. It belongs to a stallion about 6 years old and has been
selected because it is practically perfect; only the tip of the right nasal is damaged by
breakage. Most other skulls from the Hagerman quarry, including the type of Gidley's
Equus shoshonensis (here considered a synonym of ~· simplicidens) and the few skulls
from the Blanco fauna of Texas, the type locality of Equus simplicidens, are more or
less distorted. The skull LACM/CIT 210/17047 may be used for comparison with the
Pliocene and early Pleistocene species of Europe and Asia.

Fig. 1. Equus simplicidens, about 6 years old. Early Blancan (early Pliocene),
Hagerman local fauna, Glenns Ferry Formation, Idaho. LACM Z10/17047.
Condylobasallength 55Z mm.
Equus simplicidens ranges approximately from 4 to Z.5 Ma; the age of the Blanco
fauna is about 3 Ma, the Hagerman fauna about 3.4 Ma.
THE OLDEST REPRESENTA'l1VE OF~ IN EUROPE-

~ livenzovensisBajgusheva 1978
According to Samson (1975) monodactyl equids appeared in Romania in the

middle Pliocene, in faunal assemblages correlated with the well known Vialette local
fauna of the French Central Massif. This fauna belongs to the Triversa faunal unit
(Azzaroli, 1977) and Couthures and Pastre (198Z) date it between 3.0 and 3.4 Ma. If
Samson's correlations are correct, this would be the oldest record of Equus in Europe,
but his faunal lists raise some doubt; the association of Zygolophodon borsoni with
Bison, reported by Samson, is untenable.
Samson recorded Plesippus (Allohippus) major cf. euxinicus from Malusteni (the
name major is not applicable here, being homonym of Equus major De Kay 184Z) and
Plesippus (Allohippus) sp. from Berbesti Slavesti. The records are based on poorly
diagnostic remains of dentitions and the geological age of the fossils is not adequately
documented. A third record from the early middle Pliocene is based on four isolated
teeth from Beresti, referred to Equus (Equus?) simionescui Radulesco and Samson.
The teeth actually show advanced features and may possibly belong to Equus caballus,
but their fossilization is different from the rest of the fauna (Gahunia, pers. comm.);
either they were reworked, or fossils were mixed up in the collections.
Leaving aside these doubtful records, the first occurrence of Equus in Europe
and Asia has been dated to about Z.5 Ma (Lindsay et al., 1980; Bonadonna and Alberdi,
1987), and falls in the Montopoli faunal unit of western European mammalian chronol-
ogy (Azzaroli, 1977), and at the beginning of the Pinjor stage of Siwalik stratigraphy
340
(Lindsay et al., 1980; Azzaroli and Napoleone, 1982). The arrival of Equus coincided
with a significant faunal turnover; the demise of a forest assemblage characterized by
Mammut borsoni, Tapirus arvernensis, Sus minor, Ursus minimus, and the arrival of
elephant and of cervids of large size of the genus Eucladoceros (Elephant-Equus event
of Lindsay et al., 1980; see also Azzaroli, 1983).
At this time an equid of large size is represented by fragmentary skulls, denti-

tions, and limb bones of Khapry and Livenzovka in southern Russia (Gromova, 1949;
Bajgusheva, 1971, 1978) and by poorer remains of teeth and limb bones in central Italy
(Azzaroli, 1982) and eastern Spain (Bonadonna and Alberdi, 1987).
The Russian fossils were described as Equus stenonis var. major by Gromova; as
Eguus cf. bressanus by Bajgusheva in 1971, and as Eguus livenzovensis n. sp. by the
same author in 197 8. Both specific names bressanus and livenzovensis were considered
valid by Azzaroli (1982), while major is preoccupied (see above). Eguus bressanus is
more advanced than the Russian Pliocene fossils in dental features and is believed to
belong to the later part of the early Pleistocene. Boule's Eguus major represents the
same species as Viret's Eguus bressanus.
The fossils from Italy and from Spain are rather poor but do not appear to be
different from Russian fossils and may possibly represent the same species, the valid
name for which is Eguus livenzovensis. Alberdi and Bonadonna (1983) reported this
taxon as Eguus stenonis "livenzovensis"; the present author prefers to give it full
specific rank for the reasons given here.
Classifications of equids have been based largely on the characters of the denti-
tions, occasionally also on limb bones. Gromova (1949) took a broader approach, with
a detailed analysis of cranial features, but dearth of suitable fossils for the Pliocene
and early Pleistocene - along with a dearth of careful description of fossils which had
entered the collections of western Europe long ago - severely hampered her
attempt. Even now skulls of Pliocene and early Pleistocene equids are known only
from a few local faunas, and all are damaged or distorted. Restorations of the skulls
of European taxa have been attempted here; they will be used as a basis for compari-
son and for interpretation of their systematic position.
Figure 2 is an attempt of a restoration of the skull of Eguus livenzovensis. This

task was by no means easy; the type specimen, described by Bajgusheva, consists of
three unconnected fragments and its length may be reconstructed, with some approx-
imation, with the help of a jaw from the same site. The skull from Khapry, described
by Gromova, is in two disconnected fragments and is damaged in several parts; it
belongs to a not-fully-mature animal, about 4 years old according to Gromova.
Fig. 2. Eguus livenzovensis, tentative restoration of skull, from specimens from

Khapry and Livenzovka, southern Russia. Late middle Pliocene. Condylo-
basal length approximately 600 mm.
341
Table 1. Greatest lengths of limb bones.
Eguus simplicidens Eguus livenzovensis Eguus bressanus
Radius 3ZZ-344 366-385 423

Third metacarpal ZZ6-Z5Z Z56-300 286
Tibia 358-385 385-46Z
Third metatarsal Z60-Z85 Z90-330 327-345
While the samples from Italy and from Spain were dated to approximately
Z.5 Ma, the age of the Russian fossils has not been determined directly; they are
tentatively referred to the same age because of the affinities of the faunas with those
from Montopoli and from Rincon 1.
With the help of the present restoration a comparison may be attempted

between Eguus livenzovensis and its immediate predecessor in time, the North
American Eguus simplicidens. They share many primitive features: the large size, the
retention of a well developed, although shallow preorbital pit, the deep groove along
the suture of the nasals, the transversally undulated forehead, the buccinator fossa,
the strong deflection of the braincase, and the features of the dentition. The
posterior palatine foramina are in a forward position in Eguus simplicidens (Azzaroli
and Voorhies, in prep.) and in the Livenzovka skull (this is seen in a photograph of the
palate, kindly supplied by Bajgusheva to the present author); in the Kapry specimen
they seem to lie farther back, opposite the protoloph of M3/; if this is true, this is one
of the few skull features that distinguish Eguus livenzovensis from Eguus simplicidens,
and it has been observed in one specimen out of two. The preorbital pit seems slightly
less extended caudally in E. livenzovensis, but its extent is difficult to determine
exactly and Gromova's photograph is not very satisfactory.
While the size of the skull, as far as the reconstruction proposed here is reliable,
falls within the size range of Eguus simplicidens, the limbs of Eguus livenzovensis are
more elongate. The size ranges of these two taxa are distinct (see table 1).
Are Eguus simplicidens and Eguus livenzovensis really distinct species, or should
they be considered subspecies of a single species? It seems premature to attempt to
solve the question now; it may be wise to wait for more informative fossils from the
late middle Pliocene of Europe, or of Eurasia in general.
Fig. 3. Equus stehlini, tentative restoration of skull, after

specimens from the Upper Valdarno, Tuscany. Early
Pleistocene, Tasso f.u. Condylobasal length approxi-
mately 495 mm.
342
DIFFERENTIATION IN THE LATE PUOCENE AND EARLY PLEISTOCENE
Equus stenonis Cocchi 1867
In the late Pliocene and early Pleistocene Equus stenonis is richly represented
and is clearly differentiated from Equus simplicidens and, as it seems, from Equus
livenzovensis. The richest localities for the late Pliocene in western Europe are Saint
Vallier, SenElze, and Chilhac in France and La Puebla de Valverde in Spain, but the last
locality has not yielded skulls; in the early Pleistocene the Upper Valdarno
(Montevarchi group) and Olivola, in central Italy (Heintz, 1978; Prat, 1980; Boeuf,
1986; Bonadonna and Alberdi, 1987 for France and Spain; Azzaroli, 1965, 198Z; be
Giuli, 197Z for Italy). The type (lectotype) of the species, an adult male skull with
jaw, comes from the Upper Valdarno and belongs to the early Pleistocene. The other
fossils were divided into several subspecies: vireti Prat for Saint Vallier and La
Puebla de Valverde; senezensis Prat for Sen~ze; guthi Boeuf for Chilhac. They differ
from one another mainly in body size and in proportions of the limbs, but differences
are small.
Several skulls are known from the late Pliocene and early Pleistocene. All are
damaged or distorted to a greater or lesser degree. Figure 4 is an essay of a restora-
tion; in this case the task is much easier than for Equus livenzovensis and the result is
accordingly highly reliable.
A typical feature of Equus stenonis is the deep indentation of the narial notch,
which extends caudally above the third premolar. The position of the angle of the
notch relative to the teeth is subject to variation because in old individuals the tooth
row contracts, but the great elongation of the narial opening is always apparent. The
snout is slender and elongated, the diastem very long. The upper profile of the skull is
wavy; the braincase is rather small relative to the length of the face and is strongly
deflected; the forehead is transversally undulated. Equus stenonis recalls E.
livenzovensis in the characters of the forehead, the profile, and the braincase, but the
preorbital pit is poorly developed and the posterior palatine foramina are shifted
caudally, opposite the M3/ in the type skull. The dentition also shows some progres-
sive change. The enamel of the inner fossettes is richly plicated; the protocones of
Fig. 4. Equus stenonis, restoration of skull based on the lectotype and on

specimens from the Upper Valdarno and Olivola, Tuscany. Early
Pleistocene, Olivola and Tasso f.u. Condylobasal length approxi-
mately 590 mm.
343
Table z. Size ranges of metapodials.
Third Metacarpal Third Metatarsal
Equus stenonis stenonis- U. Valdarno Z36-Z50 Z73-Z87

Equ'lls stenonis stenonis - Olivola Z3Z-Z56 Z69-Z88
Equus stenonis vireti ZZQ-Z41 Z5Z-Z79
Equus stenonis senezensis Z1Z-ZZ4 Z41-Z57
Equus stenonis guthi ZZ4-Z38 Z54-Z71
Equus stenonis pamirensis ZZ1-Z5Z Z56-Z87
Equus cf. stenonis - Selvella Z40-Z53 Z70-Z98
Equus stehlini - U. Valdarno Z01-Z15 Z39-Z54
the upper cheek teeth are generally short, but occasionally more elongated protocones
occur in the nominal subspecies; in the lower molars the ectoflexids are either deep or
shallow; in the type skull they are shallow. Variation in the depth of the ectoflexids
has been observed in samples from the same local faunas: Saint Vallie.- (see pl. Z8 of
Viret, 1954), the Upper Valdarno. Although some samples are small and unsuitable for
statistical analysis, the primitive form (short protocones and deep ectoflexids) seems
predominant in all local faunas.
As stated above, the main differences among the European subspecies is in size
and proportions. The highland subspecies _!. senezensis and _!. guthi are rather small
and slender limbed; E. vireti has a large skull and short, stocky limbs; the type form,
from Tuscany, equals _!. vireti in skull size but has more elongated limbs (see table
Z). De Giuli (1987) described Equus cf. stenonis from Selvella, near the Tuscan border
of Umbria, central Italy. The fauna was referred to the later part of the early
Pleistocene and the equid is believed to be more derived than Equus stenonis stenonis
in some details: more elongate protocones in the upper cheek teeth, more elongate
limb bones, broader articular trochleas in the metapodials. The size of the limbs falls,
however, in the range of the older early Pleistocene local faunas of Matassino in the
Upper Valdarno and Olivola in northwestern Tuscany, and the details of the enamel
pattern of the cheek teeth are highly variable in Equus stenonis in general. The
Selvella equid may possibly be a distinct subspecies of Equus stenonis; pending more
detailed information a separation at the species level seems unwarranted today.
Equus stenonis followed _!. livenzovensis in time, but this does not necessarily
imply that it was a direct descendant of E. livenzovensis; the possibility should be
considered that its specialization began in North America and was brought to the Old
World by immigration.
!9!!!!_ stenonis and its Relatives in Asia

Equus stenonis, with its peculiar specialization, is not restricted to Europe.
Skulls with identical features and similar dentitions occur in at least three different
sites in China (Azzaroli, 1987), and their ages are comprised between the late Pliocene
and the early Pleistocene.
Sharapov (1986) proposed the new species Equus (Hippotigris) pamirensis, from
the late Pliocene of Kuruksaj, Tadjikistan. This is a stocky equid with rather short
limbs. Sharapov figures an upper and a lower tooth row. In the upper check teeth the
enamel is richly plicated, the protocones are short in PZ/-MZ/ and are lingually
grooved; M3/ has a long protocone. In the lower molars the ectoflexids are shallow.
Sharapov did not describe the skull of his Equus pamirensis. In the Geological
Institute of the USSR Academy of Sciences in Moscow I saw three damaged skulls
from Kuruksaj, No. 31Z0-319, 31Z0-3ZO, 31ZQ-339. They are slightly smaller than the
type of Equus stenonis but show the typical features of this species in the dentitions,
344
facial fossa, deep narial notch, and elongated snout. The condylobasal length of the
skull No. 31ZD-3ZO, the most complete of the three, is 543 mm.
The taxon E. pamirensis is distinct from the typical E. stenonis but should better
be given subspecific rank, as Equus stenonis pamirensis.
Identical features characterize the two known skulls of the Indian species Eguus
namadicus (Azzaroli, 198Z); here, however, the dentitions differ from those of Equus
stenonis in consistent features. In the upper cheek teeth the protocones are elongated
and the ectoflexids of the lower molars are shallow in all specimens the present writer
could examine. One of the two skulls of Equus namadicus, from Jhil (described as
Equus sivalensis by Hooijer, 1949) comes from the Siwaliks of the Himalayan foothills,
where Equus is reported in a range from late middle Pliocene (Z.5 Ma) to early
Pleistocene (Opdyke et al., 1979; Azzaroli and Napoleone, 198Z). The age of the Jhil
skull is not better known. The type skull of this species, figured by Falconer and
Cautley (1849), comes from the alluvial deposits of the Narbada River in Dekkan and
is of middle or late Pleistocene age.
Eguus sivalensis is represented by more incomplete skulls; it is closely related to

Equus namadicus, from which it differs mainly in its smaller size (Azzaroli, 198Z).
Equus sanmeniensis, from the Nihewan fauna of northern China, is different.

This species is represented by a nearly perfect skull of an old stallion. It is of very
large size and shares some features with Equus simplicidens: the preorbital pits, the
groove along the suture of the nasals, the wavy profile, the strongly deflected brain-
case. The posterior palatine foramina, unlike Equus simplicidens, are placed far back,
behind the M3/, but the palate may be damaged; moreover, the skull has very much
worn teeth and this may have caused a forward shift of the third molars relative to
the foramina. The narial opening does not extend caudally as in Equus stenonis and
the protocones of the upper cheek teeth are elongated. These features separate Eguus
sanmeniensis from Equus stenonis and from Equus namadicus; the skull is reminiscent
of Equus ~ now living in Africa (cf. Azzaroli, 1966), but differs in the retention
of the facial pits, of the groove along the nasal suture, and in the strong deflection of
the braincase.
~ stehlini Azzaroli 1965
This species was described by Azzaroli (1965) on specimens from the early
Pleistocene of the Upper Valdarno, Tuscany. Its distribution, in particular its absence
from the late Pliocene and earliest Pleistocene faunas such as Saint Vallier, Olivola,
and the Matassino local fauna of the Upper Valdarno (De Giuli, 197Z; Privat, 1985),
indicates that this species was confined to the later part of the early Pleistocene
(Tasso and Farneta faunas: Azzaroli, 1977, 198Z; Borselli et al., 1980; Azzaroli et al.,
198Z; Ge Giuli and Masini, 1986). Gahunia and Vekua (1979) reported the occurrence
of Eguus stehlini in Georgia, in layers immediately overlying a section representing
the Jaramillo paleomagnetic episode, i.e., younger than 0.90 Ma.
Equus stehlini is represented in the Upper Valdarno by six damaged skulls (one
with jaw), several jaws, teeth, and limb bones. Figure 3 is an essay of restoration of
the skull, which may not be very accurate because of the poor preservation of the
specimens.
Initially Azzaroli (1965) interpreted Equus stehlini as an early relative of asses

and hemiones but later studies (Azzaroli, 198Z) showed that it is very closely related
to Equus stenonis. It is of much smaller size, which is even below the values of Equus
stenonis senezensis, but shows the same specialization in the deep narial notch, the
long diastem, and the occurrence of shallow preorbital pits and a deep furrow along
the sagittal suture of the nasals. It shares with Equus stenonis several primitive
features in the dentition, such as the short protocones of the upper cheek teeth, but
differs in some apomorphic characters. Pl/ is consistently lacking; the lower incisors
(observed in one case) bear open cups. On the other hand, shallow ectoflexids never
345
occur in the lower molars. The posterior palatine foramina lie more forward than in
Equus stenonis, opposite the commissure between MZ/ and M3/, and this may be due to
the relatively large size of the upper tooth row relative to the whole skull.
The fluvio-lacustrine, early Pleistocene deposits of the Upper Valdarno cover a

wide range of time, encompassing the Olivola and Tasso faunal units of Azzaroli
(1977). While Equus stenonis and Eguus stehlini appear clearly distinct in local faunas
of this basin (E. stenonis in the Matassino clay pit near Figline, E. stehlini in the Casa
Frata fauna near Terranova), fossils collected in earlier times, without detailed strati-
graphic record, cover the whole size range from Eguus stenonis to Eguus stehlini. This
may be interpreted as evidence that the latter species was derived from the former by
a process of size reduction (Prat, 1985) and the acquisition of some apomorphic
features in the dentition.
Evidence from later faunas (Selvella, see above) shows that this process did not
affect the whole local population of Equus stenonis, which survived in the latest
Villafranchian.
~ bressanus V'IJ'et 1954
The end of the early Pleistocene (end of the late Villafranchian) was a critical
period of rapid differentiation, extinctions, and migrations, which resulted in a
dramatic faunal turnover involving all Eurasia ("end-Villafrachian event:" Azzaroli
and Napoleone, 198Z; Azzaroli, 1983; Azzaroli et al., 1988). An equid of exceedingly
large size - the largest size ever witnessed in the Old World- made its appearance
at this time. Its dental features recall Eguus stenonis but teeth are of larger size and
more derived in the plications of their enamel.
This equid is poorly represented and its geological age may be inferred on some-
what indirect evidence (Azzaroli, 1985). Viret (1954) called this equid Eguus bressanus
and based his species on an upper dentition, a lower premolar and a first phalange
from Chagny, France, which had been described by Boule under the name of Eguus
stenonis race major. To make things more complicated, there are two fossil-bearing
horizons at Chagny and the provenance of Boule's specimens is not known. Chagny n
is of Pliocene age according to Chaline (quoted in Azzaroli, 1985); this seems too old
for an equid so advanced in dental features. Chagny I has been dated to the transition
from early to middle Pleistocene and seems more likely as the source of this fossil.
Viret (1954) and Prat (1980) reported Eguus bressanus also from Seneze. In the
writer's opinion, this site has yielded two faunas of different ages (see Azzaroli et al.,
1988). The main fauna comes from a maar and is of late Pliocene age. Bout (197Z)
recorded a second fossil-bearing horizon, the slope deposits unconformably overlying
the maar. This is the possible source of few fossils that are clearly inconsistent with a
Pliocene age: Cervalces gallicus, Canis arnensis, probably also Megalovis latifrons,
and Eguus stehlini (poorly represented). Eguus bressanus could be a member of this
younger fauna.
Some upper and lower cheek teeth possibly belonging to Eguus bressanus were
reported by Azzaroli (1985) from the Upper Valdarno, in an area where the occurrence
of Pliocene sediments is to be ruled out.
A more significant sample referable to Eguus bressanus comes from East

Runton, Norfolk. The geology of this site was described by West (1980), who distin-
guished two pollen horizons: Pre-Pastonian and Pastonian; there are no younger
deposits at this site, and both Pre-Pastonian and Pastonian belong to the early Pleisto-
cene. The mammalian fauna seems in agreement with the stratigraphic partition of
West. Among the large cervids, Eucladoceros ctenoides (possibly a synonym of E.
dicranios), identified by Azzaroli (1953), may belong to the Pre-Pastonian, and this
may be the case also of Eguus stenonis, represented by teeth and limb bones;
Eucladoceros tetraceros is common in East Runton and in the Creux de Peyrolles in
the French Central Massif, dated to the latest early Pleistocene by French authors; it
has also been identified in a karst filling at Selva Vecchia near Verona, Italy, with a
346
late Villafranchian fauna in which a Galerian immigrant, Crocuta crocuta, has made
its appearance; Cervalces gallicus occurs at East Runton and Seneze, and possibly in
the Crostolo River in northern Italy, in levels overlying a late Villafranchian faunule
(Ambrosetti and Cremaschi, 1976). These two cervids may safely be assumed to be of
Pastonian age.
The age of Equus bressanus is thus subject to definite constraints. A Pliocene

age, although unlikely because of the advanced features of the dentition, is theoreti-
cally possible for the fossils from Chagny and from Seneze, but not for the sample
from East Runton. On the other hand, a dating to the earlier part of the late
Villafranchian is ruled out at Seneze and Chagny, where levels of this age have not
been reported. A latest Villafranchian age, on the other hand, is compatible with the
three localities.
The fossils from East Runton were briefly examined by the present writer but
were not fully described. Table 1 gives the main measurements of the limb bones.
The teeth are large, with a richly plicated enamel.
The early Pleistocene ended thus with an equid of gigantic size, Eguus bressanus,
and the very small Eguus stehlini. While the latter is closely related to Equus
stenonis, the affinity between this species and Eguus bressanus may be inferred from
the feature of the dentition, but is not conclusively proven.
GALERIAN EQUIDS
Equids were deeply involved in the faunal turnover which took place at. the
Villafranchian-Galerian transition, in a short time interval comprised within the span
represented by the Jaramillo paleomagnetic episode, and probably took only a part of
this span (Azzaroli, 1983; Azzaroli et al., 1988). Several species names were proposed
for equids that lived during the Galerian interval: Equus siissenbornensis Wiist 1901,
Equus altidens Von Reichenau 1915, Eguus marxi Von Reichenau 1915, Equus
mosbachensis Von Reichenau 1915, and others that will be discussed later. Most
names were based on highly unsatisfactory material. In spite of this, some at least
represent valid species, while others are not valid.
~ stissenbornensis Wiist 1901
A typical early Galerian species is Equus sussenbornensis, an equid of large size,

although not as large as Equus bressanus, and with a highly complicated enamel
pattern in its cheek teeth. After WUst, this species was described in detail by Von
Reichenau (1915) and by Musil (1969, 197Z). It occurs also in bone breccias in northern
Italy near Verona (not published), in several localities of the Forest Bed series of
Norfolk (unpublished observations by the author), and at Akhalkalaki in Georgia, USSR
(Vekua, 1987). Unfortunately, no skull has been found, so that its systematic position
has been interpreted in different ways. Musil ruled out any affinity between Equus
siissenbornensis and Eguus stenonis; at the other extreme Nobis (1971) and Samson
(1975) considered Equus siissenbornensis, Equus altidens, and Eguus marxi to be closely
related to Equus stenonis. According to Azzaroli (1985) there is no sharp cut distinc-
tion between Eguus bressanus and Eguus siissenbornensis, although the latter is less
extremely large and shows a more richly plicated enamel in the cheek teeth; Eguus
bressanus may be phylogenetically intermediate between Equus stenonis and Eguus
sUssenbornensis. A mandible from Akhalkalaki (not figured by Vekua) shows a very
long diastem, remininscent of Equus stenonis.
Measurements of the metapodials were given by Musil and by Vekua. The bones
are robust, larger than those of Equus stenonis, but do not attain the size of Eguus
bressanus.
347
~ altideas Von ReicheDau 1915
This species was based on a small sample of isolated upper and lower cheek teeth
and a complete row of lower cheek teeth from Si.issenborn, Germany. Reichenau's
nomenclature is confused. On p. ZS of his paper this species is called Equus altidens n.
sp.; on p. 149 the name has been changed to Hippotigris altidens v. Reichenau. The
name was due to the great crown height of the lower cheek teeth, whic~ does not
seem to have a counterpart in the upper teeth.
Musil described several teeth and few limb bones from the same locality. The
dentition shows some primitve features. The teeth are of moderate size, the proto-
cones of the upper cheek teeth vary from very short, as in the type of Eguus stenonis,
to moderately elongated; the enamel is never very richly plicated; the ectoflexids of
the lower molars are consistently deep, the linguaflexids are pointed, but less sharp
and more irregular in shape than in Equus stenonis.
Two metacarpal and two metatarsal bones from the same locality were tenta-
tively referred to this species by Musil. This is quite reasonable because they repre-
sent the smallest sized equid in Siissenborn, as also do the teeth referred to Eguus
altidens. The metapodials are elongate and very slender; they are equal in size and
proportions to the metapodials of Eguus hemionus.
Bones with similar size and proportions were reported by Gromova and Dubrovo
(1971) and by David and Shushpanov (197Z) from early mid-Pleistocene faunas of
Moldavia. Similarly, slender bones (metapodials and phalanges) occur at Trimingham
and at West Runton, in the Forest Bed series of Norfolk. In the metapodials the distal
portion of the volar face, above the articular keeled trochlea, is flat, like in Equus
hemionus. There is no trace of the ridge that in most other equids, including Eguus
asinus, extends upward from the keel. In the sample from Trimingham the present
writer also observed few teeth possibly belonging to Eguus altidens.
It may be concluded that an equid of medium size, with a still rather primitive
dentition and very slender limbs, populated Europe during the Galerian. In spite of the
inadequacy of the type specimens, the valid name for this equid is Equus altidens.
Musil (1969) assumed a close relationship between this species and the living Eguus
hemionus and based his assumption on the characters of the dentition. It may be
remarked that his interpretation is supported by the features of the metapodials. Only
the characters of the lower molars seem in contrast with this idea. Much emphasis
was placed by recent American authors (Skinner, 197Z; Bennett, 1980; Dalquest, 1987)
on the lower molars. Deep ectoflexids were believed to characterize zebras and
horses s. str.; shallow ectoflexids, on the other hand, would characterize hemiones and
asses. As a matter of fact, shallow ectoflexids are a common, but not strictly consis-
tent, feature of the molars of hemiones and asses (Eisenmann, 1981), and, on the other
hand, they also occur in the North American species Eguus niobrarensis Hays and
Equus lambei Hays, which in skull and skeletal features appear to be closely related to
Eguus caballus (Azzaroli and Voorhies, in prep.). It was stated above that shallow
ectoflexids occur occasionally in Eguus stenonis and consistently in Equus namadicus
and Equus sivalensis, which are not related to either hemiones or asses.
In the writer's opinion the depth of the ectoflexids may be a valid character for
distinction at species level, but shallow ectoflexids were developed independently in
separate lineages and provide no reliable basis for classification at supraspecific level,
nor for phylogenetic analysis. The hypsodonty of the lower molars, the slenderness of
the limbs - which is particularly apparent in the first phalanges - and the absence of
the ridge above the distal articular keel of the metapodials are more valid evidence
for the affinity between Equus altidens and Equus hemionus.
The present writer identified as Eguus altidens two mandibular fragments, a

juvenile palate, a metatarsal with its three phalanges, two other isolated metatarsals,
a metacarpal, and a phalange from the Lakhuti 1 local fauna of Tadjikistan; this site
lies at the base of the Jaramillo paleomagnetic episode (Dodonov, 1980; Azzaroli,
1983). This is possibly the oldest record of our species.
348
What is~ stenonis granatensis?
Ruiz Bustos (1976) reported Equus stenonis senezensis var. granatensis from
Cullar de Baza I; the fauna is of Galerian age, as evidenced by the occurrence of
Megaceros savini (referred to Praemegaceros verticornis by Ruiz Bustos), Cervus
acoronatus, Bison sp., Sus cf. scrofa, Arvicola mosbachensis, Equus aff.
siissenenbornensis. The upper dentition is vaguely reminiscent of Equus stenonis, but
bears more elongated protocones (see Pls. 16, 17 of Ruiz Bustos); in the lower
dentition the molars bear deep ectoflexids, while the shape of the linguaflexids is
rather variable (ibid., Pl. 18). Two metacarpals fall in the size range of Eguus stenonis
senezensis but are much more slender and a first anterior phalange is also very
slender. Ruiz Bustos reported a similar phalange from Lachar. Both sites are in the
Granada depression in southern Spain.
A similar equid from Venta Micena, also in the Granada depression, was
described later by Alberdi and Ruiz Bustos (1985) as Eguus stenonis granatensis n. ssp.,
and again, in more detail, by Marin (1987) from the same site. The Venta Micena
specimens do not seem to be strictly identical to those from Cullar de Baza. The
protocones of the upper cheek teeth, according to the reports and the figures, seem to
be shorter, as an average, in the Venta Micena sample; but apart from this detail the
two populations seem to be closely similar.
Agustl et al. (1987) dated Venta Micena to the later part of the early Pleisto-
cene, somewhere around 1.Z Ma. The present writer cannot agree with this interpre-
tation. The occurrence of many late Villafranchian holdovers in the fauna may be due
to the marginal geographic position of this site but the occurrence of unmistakably
Galerian immigrants, in particular Megaceros solilhacus, Bison, Soergelia, definitely
point to a Galerian age, i.e., not older than the Jaramillo paleomagnetic episode.
The systematic position of "Equus stenonis granatensis" is puzzling. The

supposed affinities with Equus stenonis are far from obvious while the slenderness of
the limbs seems rather to point to a close relationship, if not identity with Equus
altidens, and the dentition is compatible with this interpretation.
~ caballus L.
This is a well known species, which hardly needs description. It was recorded
throughout Europe in faunas ranging from Galerian to late Paleolithic and survived the
faunal crisis of the final Pleistocene in restricted areas, until it was domesticated at
the beginning of the age of metals (Azzaroli, 1985a; Telegin, 1987). During this long
time span it differentiated into a variety of morphotypes which some authors
separated as distinct species: e.g., mosbachensis Von Reichenau; steinheimensis Von
Reichenau; taubachensis Freudenberg; germanicus Nehring; abeli Antonius. The
Mongolian wild horse Eguus przewalskii has also been recorded from Europe. Gromova
rightly gave all these taxa subspecific rank within the Linnean species and added new
subspecies: e.g., chosaricus and missi. Other subspecies were added later: Eguus
caballus casariensis Altuna, Eguus caballus lenensis Romanov, Equus mosbachensis
palustris Bonifay, Eguus caballus gallicus Prat, and others still which will not be
mentioned here. The Galerian Equus marxi Von Reichenau, poorly represented by
remains of teeth, may also be a synonym of Equus caballus.
Eisenmann et al. (1985) drew attention on the occurrence of "long snouted" and
"short snouted" morphotypes, in which the different proportions are not the result of
scaling but are expression of genetic differences. The length of the snout is defined in
this case as the distance from the anterior border of the incisors to the anterior
border of the second premolars. The description as long and short snouted seems to
imply a noticeable morphological difference in the skulls, but the only apparent
difference in the cases discussed by Eisenmann is the length of the tooth rows.
Eisenmann is right in assuming that these differences are the expression of genetic
factors, and there is no wonder that such changes occurred in the 800 or 900 thousand
years that horses s. str. populated Europe. More puzzling is the fact that genetic
349
differences apparently implying evolutionary divergence dating between 130 and Z10
thousand years ago were detected in domestic horses (George and Ryder, reported by
Eisenmann et al., 1985). Horses were first domesticated in southern Russia, appar-
ently from a single stock, but possibly small wild herds survived scattered in central
Europe and in the Iberian Peninsula during the age of metals, and may have been
crossed with domestic breeds.
Pehaps the oldest record of Equus caballus in Eurasia is in the Lakhuti Z fauna of
Tadjikistan, dated about 0.9 Ma (Azzaroli, 1983). In Europe it occurs in Galerian
faunas of several sites, ranging from Moldavia through central Europe to England,
Italy, and Spain. It was very common during the late Pleistocene and declined rapidly
at the end of the Paleolithic.
Equus caballus is an immigrant from North America. It is represented by

scattered remains in Irvingtonian and Rancholabrean deposits and is very common in
the late Pleistocene to Holocene bogs of Alaska. Probably the oldest record of this
species is a mandible from the early Irvingtonian Red Cloud Formation of Nebraska
(Azzaroli and Voorhies, in prep.).
LATER MIDDLE PLEISTOCENE AND LATE PLEISTOCENE EQUIDS
As stated above, Equus caballus ranged from the beginning of the middle
Pleistocene to the Holocene. After the end of the Galerian other species occurred in
Europe and will be discussed in the present section.
The Equid from Lunel Viel Cave, France
Bonifay (1973) reported the occurrence of an equid, referred to Equus

hydruntinus, from the late middle Pleistocene cave of Lunel Viel, in southern France,
correlated to the Mindel-Riss interglacial. To the author's knowledge, this equid has
not yet been described. The author had an opportunity to see a fragmental skull and
some limb bones of this equid. The skull seems to betray some affinity with asses in
the laterally protruding posterior borders of the orbits. The protocones of the upper
cheek teeth differ from those of Equus hydruntinus in their greater elongation and in
the presence of a slight, but distinct lingual groove. The posterior palatine foramina
lie in a forward position, opposite the MZ/. The limb bones are slender, but not
extremely so as in Equus hydruntinus, and are of distinctly smaller size.
~ graziosii Azzaroli
This species was recorded in the late Pleistocene (last interglacial or more
probably beginning of the last glaciation) Maspino gravels near Arezzo, eastern
Tuscany, and from similar deposits buried under the floodplain in the Chiana valley
near Arezzo (Azzaroli, 1966, 1979). The type specimen is a fairly good, although
damaged, skull (all the occipital portion is lacking); other specimens are several jaws,
an epistropheus, a scapula, a pelvis, two radii, and a femur. Equus caballus is also
present in the same deposits and is more common than Equus graziosii. Remains of
domestic animals were also collected in the Arezzo area and were mixed with fossils
in the collections, but are easily separated by their preservation; there are few
remains of sheep, pig, two jaws of domestic ass, and a skull of a dog.
The fossils referred to in this section are well fossilized and show the same state
of preservation of rhinos, bisons, giant deer, and elephants which occur in the accom-
panying fauna.
Eguus graziosii was described in detail by the present writer (1979) and there is
little need to comment. A revision of a fossil collection in the Archaeological
Museum of Arezzo (unpublished thesis by D. Marzelli, 1980) added some specimens to
the original list.
350
The skull shows close affinities with Eguus asinus, from which it differs only in
details of the muzzle; the scapula, the pelvis, and the slender radii are also typically
asinine. However, the ectoflexids of the lower molars are deep or moderately deep in
our species, while they are generally shallow in asses, where deep ectoflexids were
observed only in few cases (Eisenmann, 1981).
~ hydruntinus Regiilia
This species was described in detail by Stehlin and Graziosi (1935) and by
Gromova (1949, 1960) on the basis of dentitions, several phalanges, bones of the carpus
and tarsus and few metapodials. Most fossils come from late Paleolithic human
settlements and this is the reason why so little is known of the rest of the skeleton,
but recently De Giuli (1983) retrieved a complete humerus and radius, some
metapodials and phalanges from a sinkhole in the Salento Peninsula in southeastern
Italy, the type area of this species. The skull is not known and this has opened room
for debate. Eguus hydruntinus was considered a relative of hemiones by Stehlin and
Grazioni and by Gromova, because of the extreme slenderness of its limbs; on the
other hand, Davis (1987) placed it among zebras because of the characters of its
dentition. The present writer is inclined to favor the former interpretation, but the
question is far from settled.
Equus hydruntinus is particularly common in the late Pleistocene of southern

Europe, especially southern Italy, including Sicily and some smaller islands. Its
skeletal characters may imply an adaptation to open, semiarid or even arid grasslands,
like the present-day hemiones. The late Pleistocene phases of low sea level favored
the formation of vast pasturages along the Mediterranean coasts, with enormous
expanses of flat areas in the Adriatic Gulf where this small equid could multiply
practically without limits. This is probably the reason of its occurrence in large
numbers particularly in the Salento Peninsula; change of habitat due to postglacial sea
rise and human overkill may have been the cause of its sudden disappearance at the
end of the Pleistocene.
CONCLUSION
With their rich and complex evolution, equids offer a valuable guide to strati-
graphic subdivision and correlation, but their identification is often difficult because
of the scarcity of well preserved skulls, which is the result of their fragility. The first
immigration of equids dates from about Z.5 Ma, with the primitive Eguus
livenzovensis. This was followed by Eguus stenonis, which may either be a descendant
of the former or an immigrant, and which survived until the end of the early Pleisto-
cene, differentiating into a variety of subspecies. At the close of the early Pleisto-
cene it presumably gave rise to the gigantic Eguus bressanus and through this, in the
Galerian, to Eguus siisf!enbornensis. With this species the lineage of Eguus stenonis
came to an end. Before Equus siissenbornensis became extinct another wave of migra-
tion brought into Europe Eguus caballus and Eguus altidens, respectively a horse s. str.
and an ass, or hemione. While Eguus caballus survived throughout the middle and late
Pleistocene, differentiating into many subspecies, the record of ass, or hemione-like
equids in Europe is much less complete. In the fossil record they appear isolated, both
in their phylogenetic histories and in their stratigraphic distribution. Eguus graziosii
is clearly related to asses, and so is possibly also the equid from Lunel Viel, while the
position of Eguus hydruntinus is less clear. One may postulate a survivial of asses or
hemiones (or alternatively of zebras) in Europe throughout the middle and late
Pleistocene, or at the opposite extreme a series of immigrations from Asia or from
Africa which would have given rise to short lived, more or less widespread coloniza-
tions.
351
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REFERENCES
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354
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355
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Cambridge University Press, Z03 p.
ADDENDUM- CHRONOLOGY
The list of species having been completed, an attempt will be made to

summarize their stratigraphic ranges. The scale of mammal units defined by Azzaroli
(1977) and developed by Azzaroli et al. (198Z, 1988) will be used here.
Equus livenzovensis falls in the Montopoli unit.
Equus stenonis has a wider range, from the Saint Vallier to the Farneta unit.
The several subspecies into which E. stenonis differentiated may offer a basis for a
more detailed stratigraphic partition. For the time being, however, such an attempt is
premature and only the nominal subspecies (Olivola and Tasso units) and the subspecies
vireti (Saint Vallier unit) seem reliable for stratigraphic analyis. It may be pointed out
that Torre and Masini (this meeting} will propose a further subdivision of the St.
Vallier unit.
Equus stehlini ranges through the Tasso and Farneta units and even higher up in
Georgia, if the identification of Gahunia and Vekua is correct.
Equus bressanus belongs to the latest Villafranchian. It occurs in association

with species that are alien to the Farneta fauna but may be correlated with some
approximation with the Farneta unit, possibly also with the following Villafranchian-
Galerian transition interval.
Equus s\issenbornensis characterizes the early Galerian and seems to be lacking

in the later Galerian.
Equus caballus ranges from Galerian to Holocene. It is surely present in the late
Galerian and possibly in the early Galerian, if the interpretation of ~· marxi as a
synonym of~· caballus is correct.
The chronologie positions of Equus graziosii and of the equid from Lunel Viel
were stated above.
Equus hydruntinus is particularly common in the latest Pleistocene. According

to Stehlin and Grazioni (1935) and to Prat (1968), it appeared during the penultimate
glaciation. The record is, however, poor and the identifications may not be fully
reliable.
356
BIOEVENTS AND MAMMAL SUCCESSIONS
IN THE SPANISH MIOCENE
S. Moya-Soia and J. Agustl
Institut de Paleontologia "M. Crusafont"

Sabadell, Spain
INTRODUCTION
In the last years, our knowledge of the Miocene mammalian faunas from Spain
has been considerably increased. Besides the presence of sequences of localities
placed in continuous sections in different continental basins (see Agustr, 1982; Daams
et al., 1981; Mein et al., 1983), there is also a good representation of large mammal
sites included in these sections. Therefore, a tentative correlation can be established
since our view of the general mammal succession seems to be rather complete. In this
paper, we will evaluate the degree of congruence between the large mammal (mainly,
artiodactyls) and small mammal (mainly rodents) records during the Miocene.
In the case of the artiodactyls, only the extinction or immigration events at the
generic level have been considered. The endemic European taxa have not been used in
our analysis (for instance, Triceromeryx, an Iberian offshoot of Palaeomeryx kaupi in
the early Aragonian; or Heteroprox, a European descendant of Procervulus). A number
of faunal groups have been established with the artiodactyls, based on immigration
events. The extinction events have not been taken into account because of their
potential diachrony among basins; for instance, Hispanomeryx disappears at the base
of the Vallesian in the Valles-Penedes basin and in the early/late Vallesian boundary in
the Duero basin.
In the case of the rodent succession, we have assembled the data from different
biozonations in the Spanish Miocene basins proposed in recent years (Van der Weerd,
1976; Agustf, 1982; Daams et al., 1981). A correlation has been established between
the artiodactyl and rodent successions and the European Mammal Neogene (MN) units
(Mein, 1975).
The obtained data have been compared with information derived from paleo-
climatic curves or paleogeographic schemes, in search of the cause of these faunal
events. Other aspects, such as the Africa-Europe interchanges and long distance
correlations have also been discussed. Finally, a general pattern of faunal succession
in the Iberian Peninsula and its correlation with Asia or Africa has been proposed.
THE ARTIODACTYL SUCCESSION
The succession of artiodactyl faunas in the Iberian Peninsula allows its grouping
into fourteen assemblages, ranging from the Ramblian to the late Turolian.

MN
I; ~!ai SPANISH
LOCALITIES
FIRST
APARITION
CORRELATION WHIT OTHER LOCALITIES
BASED IN ARTIOS
AFRICA
EURASIA
EURASIA
AFRICA I
OTHERS
EVENTS
c
,,
COI'~COLES
BUNOL
Eotragus
LA ROMIEU
Tuootiderta.
4a ' IV PAPIOL
Bunolistr iodon
EL CANYET Lagomeryx 2
8 SANT MAMET
ARTENAY ALIVEilY KAMIAL
MORATILLA
j~-
CHILLEURS
AGREDA Palataomeryx
. 3b A III Dorcatherium
MOLi CALOPA Acteocemas WINTERSOORF CHAMTWAilA
Aurelianoc~r.
WEST ?
COSTA~ NAPAK Proboscidea Oorcatherium
~
ESTilEPUY Anchitherium
Procervulus
Lagomeryx 1 LES BELLIEU~
3a
z II
v
LORANCA Teruelia JLOWER
Xenohyus BUGTI
NAVAI'lflETE Braci!Yodus Brachyodus Giraffids 1
2b VALQIJEMADO LAUGNAC
I Andegameryx :sT. GERARD
v
CEliNA I
I
I
2a I
Fig. 1. Ramblian and early Aragonian artiodactyl locality groups, correlated with
regional and general Mediterranean (MN) zones. Correlation with other
localities and general faunistic events in the Mediterranean area.
Ramblian
Three groups of faunas can be distinguished in the Ramblian stage:
LG-L We include in this group the localities of Valquemado (Loranca basin) and
Cetina de Aragon (Teruel basin). It is characterized by the first appearance of the
Bovoidea Andegameryx Ginsburg (Morales et al., 1984; Morales et al, 1986; Ginsburg
et al., 1987). A second artiodactyl present in these faunas, Amphitragulus, is an
autochthonous form related to some Oligocene elements. The two localities ascribed
to this group can be correlated with St. Gerard (France).
LG-D. This assemblage includes the localities of Navarrete del Rio (Teruel) and
Loranca del Campo (Loranca basin). It is characterized by the entry of Teruelia
adroveri (a primitive Giraffidae of the subfamily Progirafinae, see Moya-Sola, 1987),
Brachyodus (an African anthracotheriid), and the suid Xenohyus. Andegameryx
persisted from the previous group LG-I. It can be correlated with the lower part of
the Z zone of Daams and Freudenthal (1981) and, in a general way, with the MN 3a
zone from Mein (1975). Nevertheless, the locality of Navarrete was once ascribed to
the MN Zb zone (Adrover, 1978). Until now, there is no record of the upper part of the
MN 3a zone in Spain, lacking the localities with the first "Lagomeryx" ("~·" praestans)
and Procervulus, already present in the French locality of Les Beilleux (Ginsburg et
al., 1985).
LG-DL The localities of Costablanca and Moll Calopa (Valles-Penedes basin)

and Agreda and Moratilla (Teruel basin) are included in this group. The faunal compo-
sition of these localities is completely different from the previous one: Andegameryx
is the sole "surviving" artiodactyl, while Brachyodus, Xenohyus, and Teruelia dis-
appear. On the other hand, new elements in this level are Dorcatherium, Acteocemas,
Palaeomeryx, and Aureliachoerus. The localities included here can be correlated with
the "Cricetid vacuum" event, detected among the rodent faunas (zone A from Daams
et al., 1981 and zone MN 3b from Mein, 1975).
358
MN
..a[
r~ ~! SPANISH
LOCALITIES
FII'IST
APARITION
COilllELATION WHIT OTHEfl LOCALITIES
BASED IN AIHIOS
AFRICA
EURASIA f~:~~~·
OTHERS
EVENTS
LOS VALLES F.
H X C.SANTIGA
Ll.OBAfffiA!
Decemotherium
9 CAN PONSIC
I
I-- I--
IX
NOMEnEVIUA
HOST. SUP. i
HOST. INF. "Polaeotragus"
C. BAnBEAA Barberahyus
HI-
8 C. MISSEIH Austropartax
G3
=ocemas
ST. QUIIlZE Parachleuostoch
1
LA GlliVE M FORT TERNAN
,VID MASQUEFA
C. ALMIIlALL 10cerus
/ LACIESMA,A. VAL Euprox
G2 VII MANCHONES
EL BUSTE
Caprotragoides ~AN Dill NYAKAK
Hipo>danhnae
6 PARACUELLOS I H1spanomeryx PASALAR I
MABOKO
Caprotrago~'l
~
Oriqithecids
~~
.,.L~!:i~d~n- __
SANSAN I l<ubanoctoen:l
------ -
~ -E VI
PUENTEw.LEC LES FALUNS BELOM ETCHESKAYA P\iq>~ ..., ..
G"nfficls
V' TAilAZONA Micromeryx
VIEUX-COLONGE!
Hyracids
OOONELL Conohyus
TOillliJOS
4b D
v MADiliD
MOilATINES
\ALDEMOill 1o
.li ~therium
rotragocerus
GEBEL ZELTEN
1\">Selaphin
Eotragus
Fig. Z. Middle and late Aragonian and early Vallesian artiodactyllocality groups.
Aragcmian
Eight locality groups can be recognized in this stage:
LG-IV. This faunal unit is largely represented in the VaUes-Penedes (Sant

Mamet, El Canyet, Can Mas, Can Canals, Papiol, Cases de la Valenciana), Tagus
(Corcoles), and Buiiol basins. The most characteristic artiodactyls of these phases are
Eotragus and Bunolistriodon. Andegameryx and Acteocemas disappear. This group of
localities can be correlated with the zones B and C of Daams et al. (1981) and the unit
MN 4a (Mein, 1975). Hispanotherium is already present in the last faunas of this
group.
LG-V. In this group we include the localities of Valdemoros IA (Calatayud-

Daroca basin), Torrijos (Tagus basin), Madrid, and Moratines (Madrid basin). These
localities are characterized by the entry of the first Boselaphini (Miotragocerus) and
Hyotherium and the disappearance of Lagomeryx. They correspond to the middle part
of the "Hispanotherium faunas" and can be correlated with the lower part of the zones
D of Daams et al. (1981) and MN 4b of Mein (1975).
LG-VI. Several localities in the Madrid basin (O'donell, Puente de Vallecas,

Paracuellos 5) as well as the locality of Tarazona, in the Ebro basin, can be placed
here. This assemblage is characterized by the immigration of Conohyus and
Micromeryx and the disappearance of Amphitragulus. Listriodon is already present in
the locality of Paracuellos-5, substituting Bunolistriodon. It can be correlated with
the upper part of zoneD and the zones E, F, and G1 (Daams et al., 1981) and the upper
part of zone 4b, and zones MN 5 and lower part of MN 6 (e.g., Sansan, see Mein,
1975). This faunal group corresponds to the uppermost levels with Hispanotherium.
LG-vn. This phase is largely represented in the Iberian Peninsula, especially in

the Calatayud-Daroca (Manchones, Arroyo de Val), Madrid (Paracuellos-3), and Ebro
basins (La Ciesma and La Buste). Caprotragoides, Hispanomeryx, and Euprox appear
for the first time in this level, while Triceromeryx, Aureliachoerus, and Bunolistriodon
359
disappear. The microfauna from these localities allows their assignment to zone GZ
from Daams et al. (1981) and the upper part of the MN 6 zone.
LG-VJD. We include here the localities of Sant Quirze, Can Almirall, and
Masquefa, in the Valles-Penedes basin, and the fissure filling of Escobosa (the position
of the last three sites seems more doubtful due to the scarce material). In this phase
appear for the first time the taxa Protragocerus, Stephanocemas, and
Parachleuastochoerus. On the other hand, Heteroprox, Eotragus, and Caprotragoides
became extinct. This unit can be correlated with the lower part of the G3 zone of
Daams et al. (1981) and with part of the zones MN 7 and MN 8 of Mein (1975).
LG-IX. This interval is well represented in the Valles-Penedes basin, in the

localities of Can Missert, Castell de Barb~a, and Hostalets de Pierola (lower and
upper levels). The entry of Austroportax, Barberahyus, and the first "modern"
giraffids in the Iberian Peninsula ("Palaeotragus") appear in this interval. No extinc-
tion events can be referred to this interval among the artiodactyl faunas. LG-IX can
be correlated with the upper part of zone G3 and part of zone I of Daams et.al. (1981),
plus the Fahlbuschia crusafonti zone of Agusti (198Z) as well as the upper part of the
zone MN 8 and the lower part of zone MN 9 (Mein, 1975). Entry of the equid
Hipparion takes place within this unit (for instance, in the upper levels of Hostalets de
Pierola or in Nombrevilla).
Vallesian
Two main artiodactyl groups can be distinguished in the Vallesian (although some
of the localities placed in the LG-IX group were Vallesian because of the presence of
Hipparion):
LG-X. This group includes several localities in the Valles-Penedes basin (Can
Ponsic, Santiga, Can Llobateras) and the site of Valles de Fuentidueiia (Duero basin).
The sole immigrant in this zone is Decennatherium, a member of the subfamily
Sivatherinae. The genus "Palaeotragus" disappears in this phase. It can be correlated
with part of the I zone and all the H zone of Daams et al. (1981), plus the Cricetulodon
zone of Agustf (198Z) and tlie upper part of the MN 9 zone of Mein (1985).
LG-XL We include in this faunal group the localities of Viladecavalls, Trinxera

Nord Autopista (TNA), Torrent de Febulines, and Torrent de Febulines-M (TF-M), in
the Valles-Penedes basin. Besides, the locality of Masfa del Barbo, in the Teruel basin,
must be included here also. This interval records the appearance of Microstonyx,
Schizochoerus, and the cosmopolitan Boselaphini Tragoportax. On the other hand, this
is the most important extinction event in the entire Iberian Neogene, with the exit of
Hispanomeryx, Protragocerus, Miotragocerus, Listriodon, Decennatherium,
Hyotherium, and Parachleuastochoerus. The localities in this zone can be correlated
with the Progonomr{ hispanicus zone from Vander Weerd (1976) and Agusd (198Z) and
MN 10 zone of Mein 1975).
Turolian
The artiodactyl Turolian faunas can be subdivided into three mammal units:
LG-xn. This interval includes the localities of Piera (Valles-Pened~s basin), Los
Gargantones (La Celia basin), Vivero de Pinos and La Cantera (Teruel basin), and
Crevillente-Z (Alicante). This interval coincides with the entry of Birgerbohlinia and a
Giraffidae indet., besides of the first true Cervinae ("Eostylocerus pierensis"). An
anthracotheriid is also present in Crevillente-Z. Schizochoerus, Euprox, and
Austroportax are no longer present at this level. LG-X:U can be correlated with the
Parapodemus lugdunensis zone of Van der Weerd (1976) or the MN 11 zone of Mein
(1975).
LG-XDL The localities of Los Mansuetos and Concud, in the Teruel basin, and
Casa del Acero, in the Fortuna basin, belong to this interval. This interval includes
360
MN
I; .
0 SPANISH
LOCALITIES
FII"!ST
APAI'liTION
COI'll'lELATION WHIT OTHEI'l LOCALITIES
BASED IN Al'lTIOS
AFRICA
EURASIA
EURASI..\
AFRICA f
OTHmS
EVE.NTS
14
-
LA ALBEI'lCA
LIBRILLA Paracamelus
ALMENARA-M Ovibovini 811155 IGELLA 5AHABI
13 )(N V. MORO Parabos Reduncini Agniotherium
VILLASTAR-o
--------
Macae a Dvibovini
HIPIX'Pot~l.f> Bovini
ARQUILLO Hippopotamus
-
MANSUETOS
12 xm CONCUD H1spanodorcas
"Procapreolus"
f- ~·
GAI'lGANT~
PIEI'lA
11 XII LA CANTEI'lA "Eostilocerus"
VIVffiO PI NOS 81gherbolinia
CREVILLENTE Giraffinae
-·--
NIVEL MONO
T. FEBULINA~
Microstomyx
10 XI Tl'liNCH.N.AUl; Tragoportax
Sch1zochoerus Tragop:lrtax?
VI LADECAB.
Fig. 3. Late Vallesian and Turolian artiodactyllocality groups.
the latest record of Dorcatherium, Micromeryx, "Eostylocerus," and Giraffidae indet.

On the other hand, Hispanodorcas and "Procapreolus" appear in the Iberian record.
This interval can be correlated with the Parapodemus barbarae zone of Vander Weerd
(1976) and the MN 12 zone of Mein (1975).
LG-XIV. Several localities in the eastern margin of Spain can be placed in this
faunal group: Venta del Moro (Valencia), Librilla and La Alberca (Murcia), Arquillo
(Teruel), Villastar (Teruel), Casablanca-M (Castellon), etc. Most of these sites are
characterized by a rather rich and homogeneous assemblage including Parabos,
Hippopotamus, Paracamelus, and the first Ovibovini. The genera Microstonyx,
Hispanodorcas, and Procapreolus are already absent from this assemblage. This inter-
val can be correlated with the Stephanomys ramblensis zone or with the MN 13 zone
of Me in (197 5).
FLUCTUATIONS IN DIVERSITY AMONG THE AR110DACTYLS
Different curves of diversity have been constructed for the three main catego-
ries of artiodactyls (Bovoidea, Moschoidea, and Cervoidea) in order to evaluate their
different behavior.
Bovoidea
In a general way, there is a progressive increase of diversity during the

Ramblian, with a slight decrease during the early Aragonian. During the middle and
late Aragonian the diversity increases again to the early Vallesian, where the
maximum number of species is attained. An abrupt decay in the diversity took place
at the early/late Vallesian boundary (from five to two genera). There are four extinc-
tion events, a possible surviving Aragonian form, and a new cosmopolitan immigrant,
Tragoportax, at this time. After the mid Vallesian crisis, the diversity again increases
in a gradual way until the late Turolian, where a new diversity maximum is attained.
It is interesting to note that while most of the early Vallesian bovids are forms of wet
361
MN
1rtios BOVOIDEA DIVERSITY
loc
(n' genera)
I
I
I Ill
II
2b
Fig. 4. Range chart of Spanish Miocene Bovoidea and curve

of diversity (genera).
character (Boselaphini), the early and middle Turolian species display cosmopolitan or
"open-prairie" character (Antilopini, Tragoportax, Birgerbohlinia; see figure 4).
Moschoidea-cervoidea
The general pattern of these groups is almost the opposite observed in the
Bovoidea. Their diversity increases progressively from the early Aragonian and
decreases during the Turolian. There is only a small increase in the number of species
in the late Aragonian (LG-VII): five genera are present in the Aragonian in contrast to
one in the middle Turolian and two in the late Turolian (see figure 5).
362
TRAGULIDAE CEJ'"!VOIDEA DIVEnSITY
MOSCHOIDEA (n" genera)
2b
Fig. 5. Range chart of Spanish Miocene Moschoidea and

Cervoidea and curve of diversity (genera).
Suiformes
This group shows a pattern similar to that of the Bovoidea: general increase
during the Aragonian until its climax in the early Vallesian, followed by an abrupt
decrease at the early/late Vallesian boundary. In contrast to the Bovoidea, there is no
new increase of diversity after that crisis (one genus in the early and middle Turolian,
one or two genera in the late Turolian; see figure 6).
The early/late Vallesian crisis marks the boundary between two very different
faunal associations and probably corresponds to the most important faunal event of
the continental Neogene (Moya-Sola and Agustf, 1987). During the Aragonian, before
363
MN ,.,
artios
SUI FORMES DIVERSITY
(n• genera)
ll
II
2b
-
f [!
X
-60 ~ 5 2 &~
1
~
~~
~ 0
u c 0
m • c
.;,x 0 m
:r:
" "'
Fig. 6. Range chart of Spanish Suidae and curve of diversity

(genera).
this criSis, there was a highly diversified fauna of "humid" bovids and suids, while
cervids and moschids become more and more rare. After that crisis, none of these
"humid" elements persisted and only in the late Turolian was there a new increase of
the diversity of suids and "humid" bovids. Therefore, the Vallesian crisis had very
important effects in the succession of the Neogene mammalian faunas. This event has
been detected by other authors (see Figure 7). Nevertheless, it has always been asso-
ciated more or less with the Hipparion event and its effect in the Palearctic faunas.
However, the entry of this equid at 11-lZ m.y. did not have any special effect on the
existing faunas. On the other hand, the early/late Vallesian crisis occurred somewhat
later, probably at 9-10 Ma.
364
MNO DIVERSITY DAAt--15 et al.
~ BOVOIDEA HOSCHOIOEA- CERVOI OEA SUIFORMES + TEMPE~ATU~E + HUMIDITY
2b
Fig. 7. Curves of artiodactyl diversity compared with the Daams et al. (1981)
paleoclimatic curves.
THE RODENT SUCCESSION
Early Aragonian
In western Europe a very significant event during the early Aragonian is the
entry of the so-called "Miocene cricetids" (Democricetodon, Eumyarion,
Megacricetodon, Fahlbuschia, and Cricetodon) after the extinction of the last
Eucricetodon species. This event is seen as a global one, outside the Iberian
Peninsula. Nevertheless, work undertaken in some Spanish basins has revealed that
the entry of these genera was not isochronous but displayed a step-wise pattern.
Therefore, the evidence in the Calatayud-Daroca and perhaps the Valfes-Penedes
basins indicates that Democricetodon appeared before the other genera (e.g.,
Democricetodon hispanicus from Villafeliche na, zone B from Daams et al., 1981).
A second dispersal event includes the entry of Megacricetodon, Eumyarion, and

Fahlbuschia, although the last genus could be derived by local speciation from the
previous settlement of Democricetodon (e.g., the Megacricetodon primitivus zone of
Agustf, 1982 or zone C of Daams et al., 1981). A peculiar case is that of Cricetodon.
The first Cricetodon species in western Europe (Vieux-Collonges; see Mein and
Freudenthal, 1972) are found in some younger faunas belonging to the Megacricetodon
collongensis zone (zone D of Daams et al., 1981), associated with a number of
Cricetodontine taxa of mainly eastern European distribution (Deperetomys,
Lartetomys, and Collongomys). Nevertheless, most of the M. collongensis faunas in
the Spanish basins (Calatayud-Daroca, Tagus) lack Cricetodon (Alberdi et al., 1984).
Only in the Calatayud-Daroca basin, the section of Las Planas includes an association
of M. collongensis with a primitive population of Cricetodon. On the other hand, the
first abundant Cricetodon populations are in fact found associated with the more
advanced species Megacricetodon crusafonti. This datum suggests two possible inter-
pretations. In the first case, the levels sharing M. collongenesis plus Cricetodon could
365
be very badly represented in the Iberian Peninsula. This is rather improbable given the
excellent record in the Calatayud-Daroca and Tagus basins. A second hypothesis could
result from a delay between the appearance of Cricetodon, a delay not yet recorded or
recognized in other parts of western Europe.
Late Aragonian
This is a second major dispersal event among the rodent assemblages of the
Spanish Neogene. This s~cond event especially affected the glirid association, with
the entry of Muscardinus, Paraglirulus, Myoglis, Myomimus, Tempestia, and Eliomys.
However, this interval records the effective expansion of Cricetodon within the
Iberian Neogene. A further element of interest is the entry of the "modern" eomyids,
such as Eomyops. This event is best represented in the Aragonian of the Calatayud-
Daroca basin. In the Valles-Penedes basin the late Aragonian microfaunas are char-
acterized by the entry of Anomalomys, Keramidomys, and Eomuscardinus. The entry
of these elements could correspond to the same dispersal wave in the Calatayud-
Daroca basin, since the middle Aragonian is not recorded in the Valtes-Penedes basin
due to a marine transgression.
Early/Late Vallesian
Among the rodents, this is one of the main faunal turnovers of the European
Neogene. In the Valles-Penedes basin, type area of the Vallesian (see Moy'a-Sol'a and
Agustl, 1987), the transition from the early to the late Vallesian involved the
disappearance of most of the genera which characterized the Aragonian. Within the
glirids, this is the case of Bransatoglis, Myoglis, Paraglirulus, and Eomuscardinus.
Bransatoglis is a Dryomyinae of Oligocene origin which persisted during the early (Can
Martf Vell) and late Aragonian (Castell de Barbera) of the Vall~s-Pened~s basin, until
the early Vallesian (Can Llobateres). This form is absent in the late Miocene (late
Vallesian and Turolian) of Europe. Myoglis appears in the middle/late Aragonian of
Manchones (Calatayud-Daroca basin; see de Bruijn, 1966) and is a rather common
element in the late Aragonian/early Vallesian microfaunas of the Valles-Penedes
basin; e.g., Sant Quirze, Castell de Barbera, Can Llobateres, etc. It is completely
absent in the late Vallesian levels of Spain, although it persists in France in the local-
ities of Douvre and Soblay (Mein, 1984). A similar case is that of Paraglirulus, a genus
showing a stratigraphic distribution very close to that of Myoglis. Paraglirulus
werenfelsi is present in the late Aragonian beds of Sant Quirze and Castell de Barbera,
as well as in the early Vallesian of Can Llobateres (a second species, Paraglirulus aff.
lissiensis is also present in this site). This form is already absent in the late Vallesian
of the Iberian Peninsula, although it persists in the late Turolian of France (P.
lissiensis from Lissieu; see Mein, 1984). In the case of Eomuscardinus, the last record
of this taxon corresponds once more to the early Vallesian of the Valles-Penedes
basin. Therefore, the surviving glirid genera after the Vallesian crisis are those still
persisting; e.g., Muscardinus, Eliomys, Myomimus, Glis.
This pattern can also be applied readily to the eomyids, a family represented
during the Aragonian and the Vallesian by the genera Eomyops and Keramidomys.
Eomyops appears in the Iberian Peninsula during the middle-late Aragonian
(Manchones, in the Calatayud-Daroca basin) and is still present in the early Vallesian
of Can Llobateres (E. catalaunicus). Nevertheless, this species is absent in the late
Vallesian. This is also the case of Keramidomys, present in the late Aragonian (Sant
Quirze) and early Vallesian (Can Ponsic, Can Llobateres), but absent in most of the
late Vallesian localities of Spain (a tooth of Keramidomys aff. mohleri has been found
in the locality of La Bastida, in the Seu d'Urgell basin, associated with Progonomys
cattalai). Nevertheless, the persistence of this species in the Seu d'Urgell basin could
be the result of its very northern location, in the eastern margin of the Pyrenees
range). The eomyids are present in France until the late Turolian of Lissieu, persisting
in eastern Europe even during the early Pleistocene (e.g., Estramomys from Hungary).
On the other hand, the Vallesian crisis affected the Cricetids in a different
way. Most of the lineages characteristic for the Aragonian had already disappeared in
366
the very early Vallesian, before the entry of Cricetulodon (Cricetodon lavocati,
Fahlbuschia crusafonti, Megacricetodon ibericus, Democricetodon brevis nemoralis).
A second step is the extinction of the small-sized lineage of Megacricetodon minor (M.
m. debruijni) and Eumyarion, which coincides, as in the case of Glirids and Eomyids, at
the early/late Vallesian boundary. A third step, involving the exit of Anomalomys and
Rotundomys, took place at the late Vallesian/early Turolian boundary, while
Hispanomys persisted until the late Turolian. Therefore, the cricetids display a
sequential pattern of extinction or exit, very different from that of Eomyids and
Glirids. Finally, we should emphasize that some groups of micromammals, such as
insectivores and lagomorphs, were only slightly affected or unaffected by the
Vallesian crisis.
Early/Middle Turolian
The early and middle Turolian rodent faunas are basically homogeneous and
composed of three main genera of murids and two cricetids. Among the murids,
Parapodemus and Occitanomys are the possible descendants of two different branches
of Progonomys ~· cathalai and E_. hispanicus). While Occitanomys sondaari seems to
be a descendant in situ from western European populations of!:· hispanicus, the first
Parapodemus species (!:. lugdunensis) probably evolved outside this area. Together
withE_. lugdunensis, a second immigrant in western Europe is Valerymys, a large-sized
murid of probable eastern origin.
Among the cricetids, Kowalskia (?=Neocricetodon) seems very close to the

primitive populations of Cricetulodon (C. hartenbergeri), although this form is clearly
an immigrant in the early Turolian. On the other hand, the early Turolian Ruscinomys
(R. freudenthali) probably originated in situ from late Vallesian Hispanomys (H.
peralensis).
Late Turolian
This is the fourth major dispersal event during the Neogene. Among the murids,
four new genera appear: Apodemus, Stephanomys, Castillomys, and Paraethomys.
Apodemus is the descendant of advanced populations of Parapodemus ~· barbarae),
although it is a matter of discussion whether this transition took place in situ or not.
Stephanomys and Castillomys were derived from Occitanomys by increasing their
hypsodonty. Finally, Paraethomys is an eastern immigrant related to the Asiatic
genus Karnimata. Specimens belonging to a true large-sized Karnimata species appear
in the late Turolian beds of Molina de Segura-D and M. de Segura-1 (Fortuna basin,
Murcia).
The entry of Asiatic or African elements is even more accentuated in the case
of the Cricetids, with the entry of Protatera, Myocricetodon, Calomyscus (the three
are present in the fissure fillings of Salobrena and Casablanca-M), Blancomys,
Celadensia (present in the late Turolian of Arenas del Ray, Granada basin),
Pseudomeriones (present in Casablanca-M). The sole cricetids of probable central
European origin are Cricetus, a very frequent element, and Epimeriones, found in the
late Turolian of La Cerdanya basin (northern Catalonia), associated with Sminthozapus
janossy, Muscardinus aff. vireti, and Apodemus gudrunae. The sole extinctions in this
phase correspond to the murid Valerymys and the cricetid Kowalskia (replaced by
Cricetus).
LONG DISTANCE CORRELATIONS
The correlation between very distant basins or even between different conti-
nents by means of rodents is very difficult because of the possibility that vicariant
forms may occur in local areas. This is not the case with regard to the large
mammals, especially perissodactyla and artiodactyla, whose record of dispersal events
permits a more secure correlation.
367
Ramblian
Among others, the beginning of the LG-n is characterized by the entry of the
giraffid Teruelia and the African anthracotheriid Brachyodus. Teruelia is directly
related to the genus Progiraffa, a form present in the Bugti beds from Pakistan.
Therefore, a Ramblian age could be hypothesized for these deposits. The entry of
Brachyodus in Europe could also be used as an element of correlation with Africa.
Nevertheless, the recent revision of the Siwaliks anthracotheriids by Pickford (1987)
suggest their absence in other Asiatic sites. Therefore, this genus seems to have a
limited value for long distance correlations (see figure 1).
At the beginning of the LG-m one of the most important faunal events of the
Neogene occurs, with the entry of Dorcatherium, Acteocemas, Palaeomeryx, and
Aureliachoerus. Dorcatherium, recorded in Europe, Asia, and Africa, presents a wider
biogeographic distribution. If the entry of this taxon into Europe and Africa (from
Asia) was isochronous, it would be of great value for early Miocene correlations. The
earliest Miocene levels in Africa contain Dorcatherium and they can be tentatively
correlated with the late Ramblian. On the other hand, there is no clear evidence for
the presence of Palaeomeryx in Africa.
Aragcmian
The entry of Eotragus in Europe in the LG-IV unit corresponds to the first dis-
pe~sal event of the Bovids, a group of Asiatic origin. Eotragus is known from the
levels of Maboko and bmbo in Africa. Nevertheless, the presence of Listriodon in the
same beds strongly suggests that this bovid settled in Africa somewhat later than in
Europe (see figure Z).
In the early part of the LG-V unit the first Boselaphini (Miotragocerus) and the
suid Hyotherium appear. The presence of Boselaphini in Gebel Zelten (Hamilton, 1973)
suggests a middle Aragonian age for these beds.
The next artiodactyl event corresponds to the entry of Micromeryx and

Conohyus (LG-VI). The late part of this unit also records the entry of Listriodon, an
element already present in the French locality of Sansan. The locality of
Byeolometcheskaya (URSS), lacking bovids of the Caprotragoides group and Listriodon,
can be included within this group. This Russian locality also records the first entry of
Asiatic (Hypsodontinae) and African elements (Giraffidae, Kubanochoerus) in Europe.
Other African elements which enter Europe during this interval are the Pliopithecidae
and, somewhat later, the Dryopithecidae. Therefore, this interval implied a very
intensive faunal exchange between eastem Europe, Asia, and northem Africa. How-
ever, the changes were quite limited in western Europe (in fact, giraffids of the
"Palaeotragus" kind appear considerably later).
From the three elements which characterize the LG-VII (Caprotragoides,

Hispanomeryx, Euprox), Caprotragoides displays the wider geographic distribution
(Europe, Asia, and Africa). This genus seems to be of Asiatic origin, although its
oldest known record is in the Turkish localities of yandir and Pa't<llar, and Sansan in
France (see Kohler, 1987). In Africa, Caprotragotdes is known from the levels of
Nyakak (Thomas, 1984). If the dispersal of this taxon was synchronous in Europe and
Africa, Nyakak can be correlated with the base of the late Aragonian. On the other
hand, absence of Caprotragoides in Gebel Zelten suggests that this locality may be
older than Sansan. Another good chronologie marker could be Hispanomeryx, a form
usually associated to Caprotragoides. It is present in Pa~alar and 9andir as well as in
many Spanish localities belonging to zone G of Daams et al. (1981). It is, in turn,
unknown in Africa. The absence of these two taxa at Byelomechteskaya in Russia
suggests an older age for this site. On the contrary, its presence in PCifalar permits
assignment of this Turkish site to the base of the late Aragonian (MN 6) instead of its
classical assignment to the middle Aragonian (MN 5).
368
VaUesian
After the early/late Vallesian boundary, the Aragonian bovids disappear,

replaced by the cosmopolitan boselaphini Tragoportax, an element of Asiatic char-
acter. The Vallesian mammal age is poorly represented outside the Iberian
Peninsula. Therefore, there exists some difficulties in identification of the Vallesian
crisis in other areas. In Turkey, the localities of Esme-Akcacoy (early Vallesian) and
Kayadibi (late Vallesian or early Turolian) display two completely different faunal
assemblages. The bovids from Esme have clear Aragonian affinities (Protoryx
solignaci and Miotragocerus), while the fauna from Kayadibi show a completely
renewed aspect, with two species of Tragoportax and other elements of late Miocene
character (e.g., Plesiaddax, Nisidorcas, Oioceros, and Palaeoryx). In Africa, the
Ngorora formation presents a middle Miocene bovid association with Protoryx,
Homoiodorcas, Kipsigicerus, and Sivoreas. During the time of Menacer, all of these
elements were absent, replaced by Tragoportax (see figure 3).
Turolian
The LG-XIV assemblage corresponds to the last mammalian renewal of the

Miocene. The presence of many elements of African, or Asiatic character, such as
Hippopotamus, Macaca, Camelids, etc. in eastern Spain,is particularly significant, thus
strengthening the possibilities of long distance correlations (see figure 3).
MAIN MAMMALIAN EVENTS DURING TilE MIOCENE
When analyzed in detail, the main extinction/immigration events of the Miocene

of Spain can be assembled into four major phases (figure 8).
Ramblian/early Aragonian: It includes the LG-ll, m, and IV artiodactyl fauna

groups, covering the whole Ramblian and the lowermost part of the Aragonian (MN 3
and 4a of Mein, 1975, and zones Z, A, B, and C of Daams et al., 1981). During this
time there is a noticeable increase in immigrants (twelve genera of ruminants) with a
similar level of extinctions. Most of these immigrants were of Asiatic origin with
some African ones. Among the rodents, there is no special immigration event during
the Ramblian. On the contrary, during the early Aragonian the most important
micromammalian event of the early Miocene took place, with the entry of most of the
Cricetids that characterize the middle Miocene (e.g., Democricetodon,
Megacricetodon, Eumyarion, Fahlbuschia, and Neocometes). Most of these genera
seem to have an Asiatic origin (for instance, Democricetodon, Megacricetodon, or
Fahlbuschia). This phase corresponds to a rather long time span (more or less, 3 Ma).
Late Aragonian: It includes the LG-VII and LG-vm groups, including the MN 6,
7, and partially 8 zones of Mein (197 5) and the zones F and G of Daams et al. (1981).
After a period with rather low faunal turnover (LG-V and VI), there is a new increase
in the frequency of extinction/immigration events (six new large mammal immigrants
in a period covering more or less 1 m.y.). As stated, among the rodents there is an
important increase in diversity, with the first occurrence of Eomyops, Muscardinus,
Paraglirulus, Myoglis, Myomimus, and Tempestia. While the immigration/extinction
balance seems to be compensated in the case of the large mammals, no special extinc-
tion events took place during this interval.
Early/late Vallesian boundary: This event differs from the others since it seems
very restricted in duration and in most of the cases involved extinction events.
Among the artiodactyls, there are only three immigrants in comparison with ten
extinctions. The extinctions affected all the suids and most of the bovids of
Aragonian origin. The extinctions also affected the small mammals, with the exit of
the eomyids, most of the glirids that characterized the Aragonian, and the cricetids
Democricetodon, Megacricetodon, and Eumyarion. This is, perhaps, the most impor-
tant crisis in the European Miocene.
369
MN artios
lot DAAMS et al. 1981
2 groups T
XX
2b
l.Main ma.nmalian event in thelberian Peninsula.
a. African Artiodactyls in the Iberian peninsula •
.J.Other African maomals in the Iberian peninsula.
4.General Africa I Eurasia inmigrants.
S.General Eurasia / Africa inmigrants.
Fig. 8. Chart of the bioevents of the Spanish Miocene.
Late Turolian: As with the latter event, this phase is very restricted in duration
and seems clearly associated with the Messinian crisis. In the case of the large
mammals, the balance between extinctions and immigrations is compensated. In the
case of the rodents, there is a very important increase of diversi~y, which affected
more than twelve genera (cricetids and murids in most of these cases). This interval
records a maximum in the presence of African elements.
There is a certain assymetry in the duration of these four major intervals.

Therefore, while the phases A and B represent a relatively large time span, with
several dispersal events, the phases C and D correspond to rather short, punctuated
events.
370
AFRICA/EURASIA INTERCHANGES
As shown in figure 8, there is a noticeable coincidence between the main four

faunistic events detected in the Spanish Miocene and the entry of African forms in
Europe. Therefore in the interval A, during the early Ramblian, the first entry of true
giraffids took place in the Iberian Peninsula and probably also in Africa, coming from
Progiraffa of the Bugti fauna. At the same time, Brachyodus enters Eurasia (figure
10). In the late Ramblian (specifically the MN 3b zone), the Gomphotheriidae dis-
persed to Eurasia, while Dorcatherium is proably also present in Africa. In the early
Aragonian the first Dinotheriids and Tubulidentata, two forms of African origin, are
found in Spain.
During the interchange B, several forms of African distribution, such as

Giraffids, Pliopithecids, and Kubanochoerids, appear in Eurasia. This is also the case
of Dryopithecidae and Hyracoids, which enter Europe at the same time or slightly
later. On the other hand, Caprotragoides and the Hypsodontinae settled Africa at this
time. Nevertheless, none of the African elements mentioned above are found in the
Iberian Peninsula during that same time. Therefore, giraffids, dryopithecids, and
hyracoids are recorded in the very late Aragonian levels (figure 8).
There is no significant African influence in the interchange C (early/late

Vallesian boundary), although Tragoportax and Sivoreas appear in Africa following this
event.
The interval D records the maximum number of taxa of African or1g1n in the
Iberian Peninsula. Among the African large mammals recorded in Eurasia, there are
Hippopotamus, Macaca, and the Reduncini. For the first time, the Africa/Europe
interchanges also affected the rodent faunas, with the presence of elements such as
Protatera and Myocricetodon in some late Turolian deposits from the Spanish basins.
Among the Eurasian elements which enter Africa, there are some Ovibovini, Camelids,
Bovini, Carnivores, etc. (Moya-Sola et al., 1984; see figure 8).
Other than these four phases of interchange, the contacts between the African
and European continental areas are minimal. There is only the record of Eotragus and
the Boselaphini which appear in Africa during the middle Aragonian (MN 4b). On the
other hand, a direct relationship between the Iberian Peninsula and Africa can be
postulated only in the last interchange. In the A, B, and C interchanges, the dispersal
route between these areas was probably the northern Mediterranean margin via the
Near East.
CONCLUSIONS
The analysis of the artiodactyl and rodent successions in the Spanish Miocene
allows the following conclusions.
Four main dispersar events can be identified for the Miocene. These events took
place in the following intervals and events:
A -- Ramblian/early Aragonian
B - Late Aragonian
C - Early/late Vallesian boundary
D - Late Turolian
These interchanges include the sole proven African/Eurasian fa~al interchanges.
Associated with these four phases there existed also four main phases of marked
stability, without important dispersal events nor African/European contacts.
The interchanges A and B have a relatively long time span (from 3 to 1 Ma). On
the other hand, the interchanges C and D appear to correspond to phenomena of
371
small duration. In the first two interchanges, the long time span and the complex-
ity of the dispersal events strongly suggest a geodynamic (tectonic) cause. In the
case of the latter interchanges (late Vallesian and late Turolian), the immigration
and extinction events are related to short time events, possibly representing
pronounced climatic oscillations.
However, a strong correlation is found between these four interchanges and the
curves of temperature and humidity observed in the continental basins.
Because of their very wide geographic distribution, the artiodactyl taxa can be
seen as useful tools for long distance correlations. Several hypotheses of correla-
tion between the European Neogene and some African and Asiatic faunas are
proposed in the following way:
Bugti beds= early Ramblian

Songhor =late Ramblian/early Aragonian
Aliveri and Kamlial = early Aragonian
Gebel Zelten = early middle Aragonian (or MN 4b in the MN biozonation)
Byelometcheskaya = middle Aragonian (or MN 4b/5 in the MN biozonation)
Pa~alar and, probably, Maboko = late Aragonian (or MN 6 in the MN bio-
zonation)
We believe that the pattern observed in the Iberian Peninsula can also be
extended to other parts of Eurasia or Africa, since their causes should be found in
paleoclimatic or tectonic events of global extent.
REFERENCES
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continental de Navarrete del Rio (province de Teruel, Espagne). Doc. Lab. Geol.
Scien. Lyon, v. 72, p. 3-47. ~
Agust(, J., 198Z. Biozonacion del neogeno continental de Cataluna mediante roedores
(Mammalia). Acta Geol. Hisp., v. 17(1-Z), p. Z1-Z6.
Alberdi, M.T., Hoyos, M., Junco, J., Lopez Martinez, N., Morales, J., Sese, C., and
Soria, D., 1984. Biostratigraphy and sedimentary evolution of continental
Neogene in the Madrid area. Paleobiologie Continental, v. 14, p. 47-68.
Bruinj, H. de, 1966. On the mammalian fauna of the Hipparion beds in the Calatayud-
Teruel basin (Prov. Zaragoza, Spain). II A. Part ll The Gliridae (Rodentia).
Konin. Neder. Akad. Wetens., Ser. B., 69, 3, p. 1-Z1.
Daams, R. and Freudenthal, M., 1981. Aragonian: The Stage concept versus Neogene
Mammal Zones. Scripta Geologica, v. 6Z, p. 1-17.
Ginsburg, L., Huin, J., and Locher, J-P., 1985. Les artiodactyles selenodontes du
a
Miocene inferieur des Bailleux Savigne-sur-Lathan (Indre-et-Loire). Bull. Mus.
Nat. Hist. Nat. Paris, 4 ser., 7, sec. C., 4, p. Z95-303.
Ginsburg, L., Morales, J ., and Soria, D., 1987. Nouvelles faunes de grans Mammiferes
d'age Miocene inferieur dans la parte oriental du bassin du Tage (Espagne).
Consequences stratigraphiques. C. R. Acad. Sci. Paris, 305, Ser. ll, P• 6Z9-63Z.
Hamilton, W.R., 1973. The lower Miocene ruminants of the Gebel Zelten, Lybia. Bull.
Brit. Mus. Nat. Hist. (Geol.), v. Z1, p. 73-150.
Kohler, M., 1987. Boviden des t'ilrkishen Miozans. Paleont. y Evol., v. 21, p. 133-246.
Mein, P., 1984. Composition quantitative des faunes de Mammiferes du Miocene
Moyen et Superieur de la Region Lyonnaise. Paleobiologie Continental, v. 14(Z),
p. 339-346.
Mein, P. and Freudenthal, M., 1971. Une nouvelle clasification des Cricetidae
(Mammalia, Rodentia) du Tertiaire de l'Europe. Scripta Geolo~ica, v. z, p. 1-37.
Mein, P., Moissenet, E., and Adrover, R., 1983. L'extension et l'age des formations
continentales pliocenes du fosse de Teruel (Espagne). C. R. Acad. Sci. Paris, v.
Z96,p. 1603-1609.
372
Morales, J. and Soria, D., 1984. Los artiodactilos del Mioceno inferior de las cuencas
centrales de Espana. Colpa, v. 39, p. 51-59.
Morales, J ., Ginsburg, L., and Soria, D., 1986. Los Bovoidea (Artiodactyla, Mamma.)
del Mioceno inferior de Espana: Filogenia y Biogeografia. Paleont. i Evol., v.
ZO, P• Z59-Z65.
Moya Sol~, S., 1987. Los Ruminates (Cervoidea y Bovoidea, Artiodactyla, Mammalia)
del Ageniense (Mioceno inferior) de Navarrete del Rio (Teruel, Espana). Paleont.
i Evol., v. Zl, p. Z47-Z69.
Moya Sola, S. and Agustf, J., 1987. The Vallesian in the type area. Ann. Inst. Geol.
Publ., Hung., LXX, P• 93-99.
Moya Sola, S., Agustf, J ., and Pons, J ., 1984. The Mio-Pliocene insular faunas from
the west Mediterranean. Origin and distribution factors. Paleobiologie
Continental, v. 14(Z), p. 347-357.
Pickford, M., 1987. Revision dessuiformes (Artiodactyla, Mammalia) des Bugti
(Pakistan). Ann. de Paleontologie, v. 73(4), p. Z89-350.
Thomas, H., 1984. Les Girraffoidea et des Bovidae Miocenes de la formation Nyakach
(Rift Nyanza, Kenya). Palaeontographica, v. 183, p. 64-89.
Weerd, A. van der, 1976. Rodent faunas of the Mio-Pliocene continental sediments of
the Teruel-Alfambra region, Spain. Utrecht Micropal. Bull., Spec. Publ. Z, p.
1-Z17.
373
THE MIOCENE RODENT SUCCESSION IN EASTERN SPAIN:
A ZOOGEOGRAPmCAL APPRAISAL
J. Apti
Institut de Paleontologia "M. Crusafont"

Sabadell, Spain
INTRODUCTION
The first papers dealing with the paleomammalian biogeography of the Spanish
Neogene were produced by Crusafont (1954, 1958, etc.), who emphasized the endemic
character of most of the Iberian faunas. This endemic character was thought to be
caused by the effects of the Pyrenees Range, which probably acted as a biogeographic
filter, isolating the Iberian Peninsula from the rest of Europe. ·
The intensive work developed by de Bruijn (1967) and Freudenthal (1963) in the
Calatayud-Daroca basi:n partially confirmed this view, since the rodent faunas of this
basin showed a high number of endemic taxa (for instance, the glirids Armantomys,
Praearmantomys, and Tempestia).
However, the analysis of the rodent faunas from the Vallesian of the Vall~s
Penedes and other Catalonian basins revealed a rather different composition, much
closer to that of other European localities and different from that of the Calatayud-
Daroca and Duero basins (Agustf, 1978). The recognition of two biogeographic
provinces - Central and Levant - was extended by Agusti (1981) to most of the
Iberian Neogene. This subdivision of the Iberian Peninsula was lately confirmed in the
Duero basin by Alberdi et al. (1981) and by Garc1a-Moreno (1987). These two bio-
provinces, here called Ibero-Levant province and Ibero-Central province, show signifi-
cant differences in the rodent content during most of the Miocene.
The exact limits of the two bioprovinces are difficult to establish. As a common
feature, most of the basins assigned to the Ibero-Levant bioprovince were opened to
the Mediterranean Sea and were affected by transgressive phases. A number of these
basins, characterized by the presence of good exposures, have been chosen for the
present paper: Seu d'Urgell, Cerdanya, Empor!la, VaHes-Penedes, Araya, Mira,
Crevillente, Fortuna, and Guadix-Baza. Besides, the late Miocene fissure infilling of
Casablanca-M has been also taken into account. ·
THE BASINS
Seud'Urgell
Seu d'Urgell is a small intramountain basin which represents part of an assem-

blage of tectonic basins in the eastern part of the Pyrenees Range. Hartevelt (1979)
subdivided the Miocene deposits of this basin into two stratigraphic units:

(j)
® ---
Fig. 1. Map of distribution of the main eastern

Mediterranean basins in the Iberian Peninsula.
1: Seu d'Urgell, Z: Cerdanya, 3: Emporda,
4: Valles-Penedes, 5: Araya, 7: Fortuna, 8:
Guadix-Baza. The number 6 corresponds to the
Casablanca karstic complex.
A) Ballestar formation: This formation includes the classical locality of El

Firal, which furnished the first remains of a fossil pongid in Spain, Dryopithecus
fontani Lartet. Two new localities, Ballestar and Can Petit, also delievered micro-
mammal remains, with Cricetulodon hartenbergeri, Anomalomys gaillardi, Eomyops
catalaunicus, Muscardinus crusafonti, Glis vallesiensis, etc. (Agusti et al., 1979). This
association allows its assignment to the early Vallesian (Cricetulodon zone).
B) Piedra formation: Within this formation, the locality of La Bastida yielded a

micromammal association including Progonomys cathalai, Anomalomys gaillardi,
Keramidomys mohleri, and Glis vallesiensis. This assemblage suggests an early late
Vallesian age for this formation.
Cerdanya
As at Seu d'Urgell, La Cerdanya represents part of the same system of intra-
mountainous Pyrenees basins mentioned previously. The sedimentary infilling is
composed of two stratigraphic units:
A) Neogeoe lower UDit: This unit furnished in the past a high number of
mammal localities (see Agust1 and Roc a, 1987, for a revision). The faunal content
allows their assignment to the Vallesian (Estevar, Sanavastre, Llfvia, Prats i Sampsor)
or, more precisely, the late Vallesian (Das, Sta. Eugenia), in this part.
B) Neogene upper UDit: This unit lies unconformably over the last one. The
locality of Can Vilella yielded a fauna containing Epimeriones aff. austriacus,
Kowalskia cf. fahlbuschi, Apodemus gudrunae, Paraethomys sp., Sminthozapus janossyi
and Muscardinus vireti (Agustl and Roca, 1987). This assemblage suggests a late
Turolian age for this unit.
The Valles-Pene!les basin is situated to the northeast of the Iberian Peninsula. It

is filled by sediments which range from the early Miocene to the early Pliocene. The
mammal faunas from this basin are widely known from the works of Crusafont and
others. In this depression, five depositional units have been recognized (Agustf et al.,
1984):
376
Fig. z. Proposed zoogeographical units for the
European Neogene: n. = Ibero-Levant
subprovince; IC = lbero-Central subprov-
ince; WE = Western European subprovince;
CE = Central European subprovince; EE =
Eastern European subprovince; As = Asiatic
province; Af =Northern African province.
A) Basal c<mglomerate UD.it: No vertebrate remains have been found in this

unit. It is of probable early Miocene (Aquitanian) age.
B) Lower continental complex: Two biozones can be distinguished in this unit:
-Zone with Pseudodryomys ibericus: This zone was defined by de Bruijn and Van
Meurs (1967) in the locality of Ateca I. It is characterized by the presence of
Pseudodryomys ibericus associated to Peridyromys murinus and Praearmantomys (the
last genus absent in the Valles-Penedes). Other species are Ligerimys ellipticus and
Melissiodon dominans. The localities of Moll' Calopa, Costablanca-n, and Sant Andreu
de la Barca belong to this zone.
- Zone with Megacricetodon primitivus: It is characterized by the association of

Megacricetodon primitivus, Democricetodon aff. hispanicus, Pseudodryomys
simplicidens, and Glirudinus modestus. Peridyromys murinus, Ligerimys ellipticus, and
Melissiodon dominans persisted from the earlier zone. This zone records the first
appearance in the basin of Glirudinus undosus, Bransatoglis astaracensis, and
Ligerimys floancei. The localities of Can Marti Vell I and n, Sant Mamet, and El
Canyet belong to this zone.
C) Marine and transitional complex: In the Penedes and in part of the Vall~s,
the middle Miocene (late Burdigalian-Langhian) is composed of sands, lutites. and
marls of marine origin, locally associated with reef formations. At the top of this
complex, the locality of Can Almirall yielded an association including Cricetodon aff.
jotae, Fahlbuschia cf. crusafonti, Megacricetodon minor, Eumyarion medium,
Miodyromys aff. aegercii, Muscardinus hispanicus, and Miopetaurista cf. neogrivensis.
D) Upper continental complex: The sections in the Upper continental complex

are the best represented in the basin, especially in the exposures seen in the more
central and northern sectors. This complex has yielded a high number of fossil locali-
ties, ranging from the late Aragonian to the early Turolian, and are assembled into
five biozones:
377
FAMILIES cn1CETIOAE SCIURIOAE GlllliOAE
''
---- ~::~:::~
---
"' ...........
'
'
'
20 10 40 10 20 !0
CniCETIDAE SCIUrliDAE Glll'liDAE
Cricetodontinae ___ _ Sciurinae - - -- Myomiminae

~
-•
cricetidoe
Cricetinae - - Petauristinae - - Giirinoe
EillTIIJ gliridae
Eumyorion
+
---·· --- --- Dryomyinae
Anomalomys
eomyidae
~ sciuridae
muridoe
Fig. 3. The rodent succession in the Valles-Penedes basin. From left to right, the
columns indicate the following aspects: succession of the most representa-
tive localities, total number of teeth, percentages of teeth per family,
percentages within the Cricetidae, Sciuridae, and Gliridae. The column in
the left corresponds to the following sites: Early Aragonian: CM-1 = Can
Martf Vell-1, CM-ll = Can Martf Vell-ll; Late Aragonian: SQ = Sant Quirze,
CB = Castell de Barber~, HI = Hostalets de Pierola (lower levels); Early
Vallesian: HS = Hostalets de Pierola (upper levels), CP = Can Ponsic, CL =
Can Llobateres; Late Vallesian: TSA-ll = Trinxera Sud Autopista (Terrassa),
TNA = Trinxera Nord Autopista (Terrassa).
- Fahlbuschia crusafonti zone: This zone is characterized by the association of

Fahlbuschia crusafonti, Megacricetodon ibericus, ~anomys dispectus, and
Cricetodon lavocati. Other species are Anomalomys gaill i, Muscardinus bispanicus,
Myoglis meini, etc. The localities of St. Quirze, Castell de Barbera, and Hostalets de
Pierola belong to this zone. The uppermost part of the section at Hostalets de Pierola
already contain Hipparion, thus being formally Vallesian.
- Cricetulodon zone: The beds belonging to this zone are characterized by the
association of the genus Cricetulodon with Eomyops catalaunicus and other persisting
elements from the previous one (Bransatoglis astaracensis, Muscardinus hispanicus,
Paraglirulus werenfelsi, etc.). This zone contains the stratotype of the Vallesian (Can
Llobateres). The localities of Santiga and Can Ponsic also belong to this biozone.
378
- Schizochoerus zone: This zone is characterized by the exit of many of the rodent
species present in the previous biozones. This is the case, for instance, of most of the
glirid species (Muscardinus and Myomimus excepted), the species of eomyids, and two
forms of cricetids (Megacricetodon and Eumyarion). It is marked by the presence of
two species of murids (Progonomys cathalai and Progonomys hispanicus), and by the
entry of eastern immigrants (for instance, the suid Schizochoerus). This zone is repre-
sented in the eastern sector of the basin (Viladecavalls, Can Casablancas, etc.).
- Rotundomys bressanus zone: This phase is present in different localities from the
Can Jofresa-Gan Perellada section, near the town of Terrassa (Valles). Cricetulodon
d. sabadellensis from the former zone is substituted by Rotundomys bressanus. The
genera Anomalomys and Hispanomys persisted.
- Parapodemus lugdunensis zone: This zone was originally described by Van der
\Veerd (1976) in the Teruel basin. In the Valles-Penedes basin only the site of Piera
can be assigned to the Turolian. This locality has yielded the following association:
Occitanomys n. sp., Prolagus crusafonti, Hipparion mediterraneum, Chalicotherium
sp., Aceratherium incisivum, Dicerorhinus schleiermarcheri, Adcrocuta eximia,
Plioviverrops querini, Stenailurus teilhardi, Microstonyx erimanthius, Birgerbohlinia
schaubi, Tragoportax gaudryi crusafonti, "Eostylocerus" pierensis.
Occitanomys n. sp. has also been found in the locality of La Celia (Murcia),
associated to Occitanomys !JOndaari and Valerymys vireti, two species that are typical
of the lugdunensis zone.
E) Plioc:eDe caatinental unit from the Peoedea: This unit is only recognizable in
the Penedes sector. It covers unconformably the deposits of the Upper continental
unit. The locality of La Fortesa yielded some remains of Rhagapodemus
hautima ensis Prolagus michauxi, and Hipparion crassum indicating the early
Pliocene early Ruscinian).
Casablanca Karstic Complex
The Casablanca karstic complex is composed of more than seven fissure infill-
ings (Agustl and Galobart~ 1987) ranging from the late Miocene to the latest
Biharian. These sites are located in the surroundings of the town of Almenara, in the
Castellon province.
The youngest infilling of the Casablanca complex is that of Casablanca-3, which

yielded a typical late Biharian fauna with Mimomys savini, Allophaiomys chalinei,
Pliomys episcopalis, Allocricetus bursae duranciensis, Apodemus aff. sylvaticus,
Castillomys crusafonti ssp., Micromys minutus, etc.
Nevertheless, most of the sites in the Casablanca complex belong to the late
Villanyan (Casablanca-1, Casablanca-4, etc.). The most characteristic locality of this
age is Casablanca-!. It is also the sole site displaying a stratigraphic sequence.
However, the assemblage is rather homogeneous, with more than ZS speciea including
Kislangia aff. rex, Mimomys medasensis, Mimomys tornensis, Stephanomys progressus,
Apodemus aff. mystacinus, Castillomys crusafonti ssp., etc.
The oldest deposit is that of Casablanca-M, which yielded a peculiar late

TuroliaD. association with Ruscinomys sp., Cricetus kormosi, Protatera almenarensis n.
sp., Myocricetodon cf. parvus, Pseudomeriones abbreviatus, Calomyscus sp., Apodemus
gudrunae, Stephanomys ramblensis, Paraethomys miocaenicus, Occitanomys sp.,
Castillomys crusafonti gracilis, etc.
Fort'IDUL Basin
The Fortuna basin is an intramountain basin filled by marine sediments ranging

from the Tortonian I and n up to the Messinian and early Pliocene (Montenat, 1977). A
detailed study of the geology of ths basin was carried out by Santisteban (1981). In the
379
borders of the basin, the terminal Miocene pass to reef and evaporitic deposits. These
deposits were recovered by deltaic and continental bed11 which sometimes present
fossiliferous levels with mammals. Three evaporitic groups were distinguished by
Santisteban (op. cit.). The lower one overlies clays and marls of marine origin con-
taining Globigerinoides elongatus and G. extremus, indicating a Messinian age. At
least three of the main sections had delivered mammal remains (Agusd et al., 1984).
Casa del Acero sectiou: The locality of Casa del Acero yielded a mammal
association belonging to the middle Turolian (Agusti et al., 1981): Petenyiella
repenningi, Schizogalerix sp., Hispanomys adroveri, Kowalskia meini, Parapodemus
barbarae, Occitanomys adroveri, Valerymys turoliensis, Eliomys truci, Atlantoxerus
sp., etc.
La Bomera sectiou: As the Casa del Acero section, this sequence belongs to
the third evaporitic group (over the levels with G. elongatus and G. extremus). It
shows the transition from green marls of marine origin to lacustrine beds, some of
them with vertebrate remains. The fauna found in La Hornera presents the following
elements: Apodemus gudrunae, Stephanomys ramblensis, Eliomys truci, Muscardinus
vireti, Hispanomys sp.
Molina de Segura section: The lower part of this sequence is formed by green
and bluish marls with intercalations of gypsum and sands overlying the levels of the
third evaporitic group. In the upper part of this section as many as eleven fossiferous
levels yielded rodent assemblages containing Stephanomys, ramblensis, Occitanomys
sp., Paraethomys miocaenicus, Apodemus gudrunae, Hispanomys sp., Cricetus kormosi,
etc. This association indicates a late Turolian age.
Guadiz-Baza Basin
The depression of Guadix-Baza is an intramountain basin, located in the Betic

mountain range, to the northeast of the province of Granada (southern Spain). A
sequence from the late Turolian to the early Toringian (middle Pleistocene) has been
established in this basin (Agustf, 1986). The sediments of the Guadix-Baza depression
were studied by Vera (1970), who distinguished, disconformably over a basal deformed
unit of Miocene age, three main formations:
Guadis formation: Formed by detritic materials belonging to a braided river

system. They cover all the western part of the basin and the margin of the western
section.
Gorafe-Bueiago formation: Formed of chemical deposits (limestone, marls,

gypsum) with lignitic intercalations. It corresponds to a lateral extension toward the
center of the basin of the fluvial deposits of the Guadix formation. The localities of
the Gorafe section (Gorafe 1, Z, 3, 4, and 5) appear in this formation.
Baaa formation: This is the richest formation in number of fossiliferous locali-

ties, especially within the Orce-Galera-Huescar sector. Three members can be differ-
entiated in this formation. The lowermost one is composed of limestones with inter-
calations of lignitic clays and suffered some deformation. It ranges from the early
Ruscinian (Botardo section) to the late Villanyian (Fuentenueva and Alquerfa
sections). The following member (red detrital member) lies unconformably on the last
one and is formed of detrital material deposited on an alluvial plain. Final~y, the
sediments of the upper member lie conformably over the last and are composed of
lacustrine, locally evaporitic, beds deposited in the context of a very shallow lake.
The materials of this last member range from the late Villanyan (section of Orce) to
the early Toringian (section of CUllar de Baza).
The succession in the Guadix-Baza basin, with more than fifty fossiliferous
levels, permits the recognition of eight different range zones:
380
- Trilophomys castroi zone: This zone is present in the Botardo and Gorafe sections
(Baza and Gorafe-Hw~lago formations, respectively). Besides Trilophomys castroi,
other elements of this zone are Ruscinomys sp., Cricetus barrierei, Protatera sp.,
Paraethomys meini, Apodemus gorafensis, Occitanomys adroveri, Stephanomys
margaritae, Castillomys crusfonti gracilis, etc.
- Mimomys occitanus zone: As in the former case, this biozone is also present in the
Gorafe and Botardo sections. The faunal content is very similar to the Trilophomys
castroi zone, although Mimomys occitanus is already present. Other elements are
Paraethomys meini and Occitanomys brailloni. Protatera is absent while Cricetus,
represented by C. angustidens, is no longer present in the youngest localities of this
biozone (for instance, Caiiada del Castano-1).
- Kislangia cappetai zone: This zone is marked by the exit of most of the cricetid
and murid species present in the previous biozone. The arvicolids, rep:t;esented by K.
cappetai, are the dominant elements in the localities. The sites of Zujar, Galera-1,
and Cafiada de Murcia-3 belong to this zone.
- Mimomys cf. reidi zone: In this zone, K. cappetai is replaced by a rather small,
high crowned Mimomys still retaining its mimomyian structures (Mimomys cf. reidi).
Besides Kislangia, Stephanomys also disappears from the fossil record of the Guadix-
Baza basin (although both persisted in other localities of eastern Spain, for instance,
the fissure infilling of Casablanca-1).
- Mimomys cf. ostramosensis zone: This zone is characterized by an abrupt diversi-

fication of the arvicolid assemblages, now represented by three different lineages:
Mimomys cf. ostramosensis, Mimomys bland, and Allophaiomys n. sp. The levels
containing this kind of fauna are located immediately below those already displaying
Allophaiomys pliocaenicus. The Murid representation is restricted to Apodemus aff.
mystacinus, Apodemus aff. sylvaticus, and Castillomys crusafonti ssp.
- Allophaiomys pliocaenicus zone: Allophaiomys pliocaenicus is represented in the

Guadix-Baza basin by populations of large size, very close to Allophaiomys ruffoi
Pasa. No species of Mimomys is present in this biozone. The localities of Venta
Micena-1 and Z (with a rich large mammal assemblages), Barranco Loon,
Fuentenueva-Z, etc. belong to this level.
- Mimomys savini zone: The most significant locality of this biozone is Loma
Quemada-1, which presents the following association: Mimomys savini, Allophaiomys
cf. burgondiae, Allophaiomys cf. nutiensis, Apodemus sylvaticus, Micromys minutus,
Castillomys crusafonti ssp., etc. The dominant arvicolid is again Mimomys, repre-
sented by M. savini.
- Arvicola cantiana zone: This zone is only represented in the Guadix-Baza. basin by
the locality of Collar de Baza. It is characterized by a typical Cromerian association
including Arvicola cantiana, Microtus brecciensis, Allocricetus bursae duraaciensis,
etc. (Ruiz and Michaux, 1976). Castillomys is no longer present. -
THE TAXA
Myomimiaae Daams
This subfamily was created by Daams (1981) assembling glirids with very simple
dental pattem such as the extant species Myomimus personatus. Its members probably
were adapted to dry environmental conditions.
Members of this subfamily characterize the earliest Aragonian (or late

Ramblian, according to Daams et al., 1987) associations of the Valles-Penedes basin.
Thus Peridyromys murinus and a form close to Pseudodryomys ibericus are the
dominant elements in the oldest fossiliferous beds of this basin (Molf Calopa,
Costablanca-n, Sant Andreu de la Barca).
381
Table 1.
t.mARND~LV~I
EARLY A1WD1LV1
primiti'fua crusafonti
VAIJ.ESIAII
.,_... llicoaal (~. 1'lBZ)
01-I CII-II SQ CB HI HS CP CL TSA-2 TNA Localities.

228 115 91 254 306 232 307 750 52 73 Total n..ber of teeth
1 Bumfarion weinfurteri
4 16 De.ocricetodon aff • .l!!!l!!!!!9!!
10 8 Heaacricetodon •· l!!!!!,tiVIB
..
2 5 Mesacricetodon 11. 11inor
n
4 46 Ewararion ~~edium
4
29
+
13 4
An0118lc.ays pudrxi
Cricetodon lavocati
...
n
..
1 26 19 Mesacricetodon ibericus
tool
32 + 6 + Fahlbuschia crusafonti
10
11
Hispan••• aff. ~
Democricetodon - . l i s
...
47
+
!J)
13
Hispanoons dispectus
Cricetulodon lmt!onberFi
..."'
tool
2 1 Me1!!5ricetodon 11. deliruiini
66 5 15 1 HisR!!!OII!S thaleri
60 Cricetulodon ....-u.n.ls
+ RotundOIIJS cf. lllllt1srotllld
16 Eum~:arion leemani
2 4 An01111l!!!IS &&illardi
46 74 Rotund011ys bressanus
8 4 Rotund011zs sp.
I
+ Proaon011ys cathalai
4 Prosonooiis aff. cathalai
10 1 ProsonomJ:s hisl!!nicus
4 27 Pseudodrzomys si!!R)idden!l
1 2 PeridtrOIIYS llllrinus
,..
C'>
3 2 Microdyr011ts sp.
...
.
1 + Glirudinus JtOdestus
3 3 + G. undosus
+
5
2
1
+ +
+
Bransatoalis astarac:ensis
Mzoslia lll!ini
...
5 5
7
3 3
3
Muscardinua hispanicus
Eolluacardinus val.lalilnds
..."'
tool
2 1 Psraalirulus .....te1si
3 1 MiodiiostS la&b:tld
1 Muscardinus heintzi
I
61 36 Liaeri•zs ellipticus
10 7 Liaeriazs florancei
1 1 lerllllid•zs canathicus
1 Eoii!02S catalaunicus
2 K.eramid!!!!s l!!rtesunatoi
+ + 11
4
4
5
7
1
24 13
2
2
2
13 20 Sciurinae
Petaurist inae
I
382
Peridyromys murinus has a very wide geographical and biostratigraphical distri-
bution, from the late Oligocene beds of the Ebro basin (northeastern Spain, see Agusti
et al., 1987) to the middle Miocene of Vieux-Collonges (France). ~is speci~s is also
present in the levels with Megacricetodon primitivus of the Valles-Penedes basin.
Unfortunately, the presence of a transgressive phase in the Valles-Pened.es does not
permit a determination of the upper limit of this species in the lbero-Levant
province. Nevertheless, Peridyromys murinus is absent in the levels with
Megacricetodon collongensis of the Calatayud-Daroca and Tagus basins (Daams et al.,
op. cit.; Alberdi et al., 1984). The last record of this species is found in the middle
Aragonian of central Europe (see Ziegler and Fahlbusch, 1986). P. murinus is con-
sidered to have had a mainly western European distribution.
As stated, Pseudodryomys ibericus appears associated to the former species in

the lowermost levels of the V~les-Pened.es and Araya basins. This species is substi-
tuted in the next biozone (primitivus zone) by P. simplicidens (although some ibericus
morphotypes are still found). Curiously, P. ibericus persists during the middle
Aragonian in central Europe (southern Germany, see Mayr, 1979). As Peridyromdii
murinus, the Pseudodryomys group displays an European biogeographic character.
the other hand, no member of the Armantomys-Praearmantomys group has ever been
found in the Valles-Penedes basin. In the Ibero-Levant province, only the locality of
Bui'lol delivered some teeth of this endemic lineage. On the other hand, Armantomys
is very frequent in the Megacricetodon collongensis levels of central Spain (Calatayud-
Daroca, Duero, Tagus) and persisted in these basins until the late Aragonian (for
instance, Armantomys tricristatus from Escobosa; see Sese, 1980).
The Myomiminae display a peculiar biostratigraphic distribution in the levels

above the transgressive phase of the VaUes-Penedes. Thus, the members of this
family are lacking in the highly diversified faunas of St. Quirze and Castell de
Barbera, where Glirinae and Dryominae have a good representation. On the other
hand, Miodyromys hamadryas appears in the poorly diversified faunas of the early
Vallesian such as Hostalets de Pierola (Hipparion levels) and Can Ponsic, and is later
absent from the fauna of Can Llobateres. According to de Bruijn (1967), M.
hamadryas seems to be in the origin of the Myomimus lineage, since Myomimus dehmi
nombrevillae from Nombrevilla is indistinguishable from the former species. If this
relationship is confirmed, most of the species now included in Miodyromys would in
fact belong to Myomimus. Myomimus dehmi still occurs in the late Vallesian of Riu
Ripoll (Valles-Penedes basin). In the levels younger than the late Vallesian, the
Myomiminae are absent from the fossil record of the lbero-Levant province. In fact, a
sort of mutual exclusion seems to exist between the Myomiminae and the Dryominae,
even in the present faunas. Thus, the Aragonian-Vallesian Myomiminae can coexist
with the Glirinae (for instance, Muscardinus), whereas they are absent in the assem-
blages where elements like Bransatoglis, Paraglirulus, or Eliomys are present. During
the Turolian, Myomimus persisted in the eastern Mediterranean, while Eliomys char-
acterized the western Mediterranean microfaunas.
Glirinae
The first Glirinae in the V~les-Pened.es basin are found in the localities of Can
Martf Vell I and n (Megacricetodon primitivus zone). They belong to the genus
Glirudinus, which at this level is represented by two different species: G. modestus
and G. undosus. Both species are widely distributed in western and central Europe. G.
modestus is also present in the basins from central Spain, while G. undosus is absent.
The former species persisted in the V~les-Penedes basin until the late Aragonian at
Castell de Barbera, being absent in the Vallesian levels.
The late Aragonian is characterized by a high diversification of the Glirinae.

Thus, besides Glirudinus, Myoglis, Muscardinus, and Eomuscardinus are found in differ-
ent localities of the Catalonian basins: Sant Quirze, Can Simeo, Castell de Barbera,
and others. This situation persisted during the early Vallesian (Can Ponsic, Can
Llobateras, Ballestar, Can Petit, etc.). However, the transition to the late Vallesian
involved a dramatic decrease of the Glirinae, which remain restricted to the genera
Glis and Muscardinus.
383
The first representatives of the genus Glis in Spain were recorded in the very
early Miocene of Cetina de Aragon (Glis tru;;lsi Daams). After this record, Glis is
absent from the Spanish Neogene until the early Vallesian. In this age, Glis
vallesiensis Agusti is recorded from the localities of Ballestar and Can Petit, in the
small Pyrenees basin of Seu d'Urgell. Glis vallesiensis is also present in the late
Aragonian of Poland (Glis sp. from Opole, see Kowalski, 1967). In the present time,
Glis is present in the Iberian Peninsula only in the northeastern corner, associated with
wet mountainous habitats. I think that this environmental constraint can be extrap-
olated to the fossil species as a way to explain their peculiar distribution. In such a
case, the presence of Glis would indicate wet and cold environmental conditions.
In the case of Muscardinus, the first true representatives appear in· the late
Aragonian (Muscardinus hispanicus from Sant Quirze and Castell de Barbera). Very
rare specimens have been quoted also from Anwil (Switzerland) and from La Grive
(France), but, in general, this genus is absent from the late Aragonian sites of central
Europe, its European expansion taking place in the early Vallesian. There are some
doubts about the origin of this taxon, although "Muscardinus" thaleri from the middle
Aragonian of Calatayud-Daroca fits well as a possible ancestor. Nevertheless, no
intermediate populations between the former species and Muscardinus hispanicus have
been described. After the late Vallesian crisis, Muscardinus reappeared in a punctu-
ated fashion in the late Vallesian (Can Jofresa, Trinxera Nord Autopista level, in the
Valles-Penedes), late Turolian (Can Vilella, in the Cerdanya basin), and early Pliocene
(Canada del Castaiio, in the Guadix-Baza basin) of the lbero-Levant province. A late
Miocene insular endemism, Muscardinus cyclopaeus Agusti, Moya-Sola, and Pons-Moya
has been described in the island of Menorca (Balearics).
Dryomyinae
The first Dryomyinae appear at a very early time in the Iberian peninsula:
Microdyromys praemurinus in Montalban and Bransatoglis n. sp. in Calaf, both of
middle Oligocene age.
In the early Aragonian, the sole members of this family are Bransatoglis and
Microdyromys. Microdyromys is very well represented in the lbero-Central province,
but is a very rare element in the Valles-Penedes basin. The .sole quotation of this
taxon is Microdyromys sp. (legidensis-koegniswaldi group) from the early Aragonian of
Can Mar:t1 Veil I and II.
Bransatoglis astaracensis is also a rare species in the localities where it is

present. Nevertheless, this taxon displays a noticeable time span in the Valles-
Penedes basin, from the early Aragonian of Can Martf Veil to the early Vallesian of
Can Llobateres. Bransatoglis is almost completely absent from central Spain (some
rare finds in the early Ramblian of Ramblar-1, see Daams et al., 1987). On the other
hand, this taxon has a wide distribution in western and central Europe.
Finally, Paraglirulus is very rarely found in central Spain, as Bransatoglis, (some

teeth in the middle Aragonian of Valhondo-4, in the Calatayud-Daroca basin, see
Daams et al., op. cit.). On the other hand, P. werenfelsi is present in the late
Aragonian (Castell de Barb~ra) and early Vallesian (Can Llobateres) of the Valles-
Penedes basin. A second species, Paraglirulus aff. lissiensis appears also in the former
locality. Paraglirulus has a mainly European distribution during most of the
Neogene. As in the case of most of the middle Miocene Glirinae, Bransatoglis and
Paraglirulus disappear at the early/late Vallesian boundary. Nevertheless, Paraglirulus
persisted in other European basins during the rest of the Miocene.
The sole Dryomyinae present in the late Neogene associations of eastern Spain is
Eliomys. The most common species during the Turolian is Eliomys truci, present in
Casa del Acero, Crevillente-6, Venta del Moro, Librilla, Salobrena, Casablanca-M,
etc. The first Eliomys intermedius appear in the Guadix-Baza basin in the very early
Pliocene levels of the Botardo section (see Agusti, 1986). Eliomys intermedius
persisted in this basin until the early Pleistocene (localities of Venta Micena-Z and
Orce-3, in the Guadix-Baza basin, see Agustf et al., 1987).
384
Cricetodontinae
The first Cricetodontinae in the Iberian Peninsula belong to the species

Megacricetodon minor primitivus. Curiously, this species, considered as the most
primitive member-or-the genus, has been only recorded in southern Europe (Valles-
Penedes and Calatayud-Daroca basins in Spain and Aliveri in Greece, see Agusti, 1981;
Freudenthal, 1963, and Meulen and de Bruijn, 198Z), while the record of
Megacricetodon in France and Germany starts with the somewhat younger chronosub-
species M. minor collongensis. In Spain, M. minor primitivus appears to be associated
with a wet environment (high diversity of glirids), while the localities with M. m.
collongensis seem to indicate more dry conditions (low diversity of glirids, represented.
by the Armantomys group). The Megacricetodon minor lineage persisted until the
early Vallesian (M. minor debruijni from Can Llobateres) in very low percentages and
is always associated with wet assemblages (high proportion of flying squirrels and
castorids).
A second Megacricetodon lineage, that of Megacricetodon crusafonti-M.

ibericus, appears in the middle-late Aragonian beds of the Iberian Peninsula. This
second lineage is characterized by a net subdivision of the anteroconid in the lower
M1. Moreover, a clear exclusion between Megacricetodon ibericus and the Cricetinae
exists in the late Aragonian-early Vallesian beds of the Valles-Penedes basin, as it is
shown in Table z. Megacricetodon ibericus is absent or very poorly represented in the
localities with highly diversified Cricetinae, Gliridae, and flying squirrels, indicating
wet biotopes. On the contrary, they are a common element in those localities lacking
Glirinae and displaying high percentages of ground squirrels (see figure 3). Curiously,
the Megacricetodon minor lineage is usually present in the wet localities. In any case,
no Megacricetodon species continues after the early/late Vallesian crisis.
The second main Cricetodontinae lineage, that of Cricetodon-Hispanomys, has a

very different history. The first member of this lineage entered western Europe in the
upper levels with MeJ:acricetodon collongensis, probably associated to a more diversi-
fied fauna of criceti s of eastern origin such as Lartetomys, Deperetomys, and, possi-
bly, Anomalomys (as it is shown in the French locality of Vieux-Collonges, see Mein
and Freudenthal, 1981). In the Valles-Penedes basin this episode is undocumented
because of the Langhian transgression. The first continental beds after this marine
event already present a form close to Cricetodon jotae or C. aquirrei. In the late
Aragonian, the tribe Cricetodontini is represented by two different forms. On one
hand, Cricetodon (Pararuscinomys) lavocati from Sant Quirze and both levels at
Hostalets de Pierola is the last representative of the C. sansaniensis-C. albanensis
lineage, which finally attains the same characteristics asthe Hispanomyslineage. C.
lavocati is no longer present in the levels with Cricetulodon.
On the other hand, a true Hispanomys species, H. dispectus, is found together

with the former in the latest Aragonian and very early Vallesian levels. H. dispectus
froll} Hostalets de Pierola seems closely related to C. aquirrei from Escobosa (see
Sese, 1980). During the late Aragonian and the whole Vallesian, Hispanomys shows a
striking diversification at the specific level, each basin displaying its own vicariant
form. This is the case of the Rhone (H. decedens from la Grive), Valles-Pened~s (H.
dispectus) and Calatayud-Daroca basinSlH. nombrevillae and H. aragoniensis). At the
end of the Vallesian, at least three different lineages of Hispanomys persisted: H.
peralensis in the lbero-Central basins, H. thaleri in the Valfes-Penedes, and H.
mediterraneus in Montredon (France). Only one of these lineages, that of H.
peralensis, survived in the early Turolian, increasing in size and hypsodonty, thus
giving place to the first species of the genus Ruscinomys, R. schaubi from Los
Mansuetos (Teruel basin). Nevertheless, an endemic form of Hispanomys, H. adroveri
from Casa del Acero (Fortuna basin) persisted in the eastern basins until the late
Turolian. On the other hand, Ruscinomys, with more progressive chronospecies
(Ruscinomys lasallei, R. europaeus) is still present in the early and middle Pliocene.
The expansion of the arvicolids ended with this lineage whose origin can be traced
back to the early Miocene.
385
CricetiDae
The first Cricetinae in Europe belong to the species Democricetodon hispanicus

present in the early Aragonian of the Calatayud-Daroca and Valles-Penedes basins.
During the first part of the Aragonian, this genus has a very wide distribution in
Europe, including Greece and Turkey (see de Bruijn and Meulen, 1979; Unay and de
Bruijn, 1984). Probably Democricetodon (as Copemys in North America) appeared as a
vicariant reply of the Asiatic taxon Spanocricetodon, in the course of an expansion of
the latter genus.
In the early Aragonian of the Valles-Penedes, Democricetodon aff. hispanicus is

a relatively abundant form (primitivus zone). This species was originally described in
the locality of Villafeliche ZA (Freudenthal, 1968) and seems so far restricted to the
Iberian Peninsula. It probably corresponds to an Iberian reply of D. franconicus from
Erkertshofen. During the middle Aragonian, Democricetodon disappears from the
Spanish fossil record, its evolution being restricted to central and eastern Europe.
A second entry of Democricetodon in the Iberian Peninsula took place in the late
Aragonian of Castell de Barbera (Valles-Penedes basin). This latter population was
described as a new subspecies of Democricetodon, D. brevis nemoralis, displaying
more progressive characters than the type population of D. brevis. Thus, the first
lower molar shows an already subdivided anteroconid, a character which is also
present in the genus Cricetulodon. This feature is also observed in advanced popula-
tions of D. gaillardi. Nevertheless, the shape of the lower molars does not support the
idea of a direct phylogenetic relationship of these species with the former genus.
Democricetodon is very rarely found in the late Aragonian of the central basins (5% of
D. aff. gaillardi in Las Planas SK, see Daams et al., op. cit.). Nevertheless, during the
middle late Aragonian, Democricetodon reached a very wide distribution, from Spain
to Greece (although not Turkey, see Unay and de Bruijn, op. cit.). We do not consider
D. kohatensis from the Chinji formation as a true Democricetodon species (see
Wessels et al., 1982). In fact, this species seems much more related to the Asiatic
Spanocricetodon stock (for instance, i· khani from the Murree formation, see de Bruijn
et al., 1981). Nevertheless, Democricetodon seems really present in Asia in the
Aragonian of Songhinzhuang (Xiacaowan, China, see Li Chuankuei et al, 1984).
The early Vallesian is characterized by the expansion of Cricetulodon, although a

first entry of this taxon is recorded in the pre-Hipparion beds of Hostalets de
Pierola. Cricetulodon has a very wide distribution, ranging from the Iberian Peninsula
to Greece. The most common species of Cricetulodon is C. hartenbergei, a species
retaining long mesolophs and mesolophids which is found both in central and eastern
Spain. The type species of this genus, C. sabadellensis, has been recorded in the type-
Table z. Mutual exclusion between the Cricetinae and the species Megacricetodon
ibericus in the late Aragonian/early Vallesian sites of the Valles-Penedes
basin. In the case of Castell de Barbera, the Cricetinae form is not
Cricetulodon but Democricetodon bi:evis nemoralis.
Cricetulodon Megacricetodon ibericus
Lower Hostalets 1% Z6%

Castell de Barbera 11%
Upper Hostalets 19%
Can Ponsic 13%
Can Llobateres 60%
386
locality Can Llobateres (Valles-Penedes basin). Since the early Vallesian record is
very scarce in the rest of Europe, we do not know the exact extent of Cricetulodon
northward. Nevertheless, the presence of some derived taxa such as Rotundomys in
France and Microtocricetus in Germany strongly suggests that this genus was probably
also present. Both Rotundomys and Microtocricetus display molars with a flat wear
surface, a character that is observed for the first time in some teeth of the Can
Llobateres population (Rotundomys sp. in Agustl, 1981; it is a matter of discussion
whether these teeth represent a real different species or belong to Cricetulodon
specimens having changed their wear habits). Rotundomys had a western European
distribution. However, in spite of some scarce finds in Portugal and the Duero basin
(Antunes and Mein, 1979; Garcia-Moreno, 1987), this genus was absent in the late
Vallesian of the Ibero-Central basins (for instance, Teruel-Alfambra, Van der Weerd,
1976). Otherwise, Rotundomys is the major component of the latest Vallesian faunas
of the Valles-Penedes and Rhone basins (see Agustl, 1981 and Mein, 1984).
Rotundomys will be replaced in the early Turolian by the cosmopolitan genus
Kowalskia (or Neocricetodon). Kowalskia strongly resembles Cricetulodon
hartenbergeri, while it is easily distinguishable from C. sabadellensis. Some isolated
populations of Cricetulodon hartenbergeri could have survived in eastern Europe or
western Asia, entering western Europe in the early Turolian. The last populations of
Kowalskia in the Ibero-Levant basins show some signs of endemism (Kowalskia
adroveri from Casa del Acero, in Fortuna, and Kowalskia sp. from Crevillente-5, see
de Bruijn et al., 1975). In the late Turolian, Kowalskia is replaced by the Cricetus
kormosi-Cricetus barrierei lineage, another cosmopolitan element. However, it must
be pointed out that the late Miocene-early Pliocene Cricetinae in the Spanish basins
never reached the degree of diversity attained in eastern Europe.
Myocricetodontinae
The first Myocricetodontinae in Spain were described by Aguilar et al. (1984) in

the fissure infilling of Salobreiia (Granada). This locality is located on the Granada
coast, in the extreme south of Spain. Nevertheless, the presence of Myocricetodon in
the Ibero-Levant province, in a more northern position is fully confirmed by the finds
of Casablanca-M (Myocricetodon cf. parvus). Myocricetodon cf. parvus from
Casablanca-M displays upper M1 in which the dominant morphotype is XZY3Z1 (sensu
Jaeger, 1977), with double anterocone, presence of longitudinal spur, and absence of
lingual cuspule between the protocone and the hypocone (although a lingual cingulum
is always present). A small cuspule is also observed in some cases between the two
lobes of the an.terocone. The Casablanca-M species seems close to the XZYZZ1
morphotypes of Myocricetodon parvus from Beni Mella! and Pataniak-6, in the
Maghreb area (Jaeger, 1977), not having attained the evolutionary degree of M. seboui
(presence of a longitudinal spur and lingual cuspule between the protocone and
hypocone). The teeth in the Salobreiia infilling can be referred to the same species as
the Casablanca-M population, that is, Myocricetodon cf. parvus.
We also consider the genus Calomyscus as belonging to the Myocricetodontinae.

The locality of Casablanca-M has yielded some teeth referable to this genus. By its
size, these molars seem to be intermediate between those of C. minor from Maritsa
(Rhodes, see de Bruijn et al., 1970) and C. delicatus from Salobreiia (Aguilar et al.,
1984). For this reason, we refer to this material as Calomyscus sp.
Gerbillidae
The first gerbils in the Spanish Neogene were quoted by de Bruijn (197 4) in the
late Turolian and early Ruscinian of Caravaca (Murcia) and Gorafe-1 (Guadix-Baza
basin, Granada). Lately, the genus Protatera has been mentioned in different late
Turolian localities in the Ibero-Levant province (Alcoy, Salobrena, and others). Never-
theless, the material is usually very scarce, not allowing an accurate description. On
the other hand, Protatera is a relatively frequent species in the late Turolian fissure
infilling of Casablanca-M. The population from this site does not fit in any of the
described species of Protatera. Therefore, we refer the Casablanca-M form to a new
species, P. almenarensis which is described below.
387
2
~]
3 4
5 6 7
Fig. 4. Protatera almenarensis n. sp. from
Casablanca-M. 1) Left M1/, ACS-M-
11. Z) Right M/1, ACS-M-33. 3)
Right MZ/, ACS-M-06. 4) Left MZ/,
ACS-M-4Z. 5) Right M/3, ACS-M-
37. 6) Right M/Z, ACS-M-03. 7)
Right M3/, ACS-M-OZ.
Plate 1. Protatera almenarensis n. sp. (Casablanca-M). 1) Left M1/,

ACS-M-07. Z) Left M/1, ACS-M-15 (Holotype).
Protatera almenarensis n. sp.
Derivatio nominis: After Almenara (Castellon), the town where Casablanca-M is

placed.
Type-locality: Casablanca-M (Almenara, Castellon).
388
Reference level: Late Turolian.
Holotype: Lower MI, ACS-M-I5, deposited in the Institut de Paleontologia

"M. Crusafont" (Sabadell, Spain).
Diagnosis: Species of Protatera with flat or concave anterior side of the

anterocone in the upper M I and very broad anteroconid in the
lower MI. M3 strongly reduced.
Differential diagnosis:
Protatera almenarensis differs from the type-species of the genus, P.

algeriensis, by its triangular shaped anterocone and by the degree of reduction of the
upper and lower M3 (see table 3).
This species also differs from P. kabulense from the late Miocene of Pul-e
Charki (Sen, I983) because of the shape of the first lower MI, much more elongated,
and by its very broad anteroconid.
Finally, P. almenarensis differs from the extant Tatera indica since this species
still retains the posterolophs in the upper molars and the M3 are not so much reduced.
Measurements: see Table 3.
Description:
M1/: The anterocone is more or less triangular, its anterior side being flat or
even concave in most of the specimens. A small enamel ring within this is observed in
one specimen. In some specimens, a posterior spur connecting the anterocone with the
protocone/paracone complex is present. This kind of longitudinal spur does not exist
between the protocone/paracone and the metacone/hypocone complexes.
MZ/: This tooth follows the morphology seen in the upper MI. Nevertheless,
the medial constriction between the labial and lingual cusps is less developed than in
the MI.
M3/: The upper M3 is strongly reduced, with a transverse crest (protocone/

paracone complex) and an extremely reduced metacone/hypocone complex, which
quickly disappears with wear.
M/I: This tooth is characterized by a very broad anteroconid with a labial

anterolophid attaining the same size as the anteroconid. A very low longitudinal spur
is present between the transverse ridges in the base of the sinusids. The posterolophid
is very reduced.
Table 3. Measurements (in mm) of the teeth of Protatera almenarensis n. sp. from
Casablanca-M (Almenara, eastern Spain).
Length Width
min. X max. min. X max. N
MI/ 2.85 2.93 3.09 2.00 2.10 2.22 3/6

MZ/ I.71 I.75 I.83 I.86 1.97 2.06 4
M3/ 1.0I 1.11 1.19 1.39 1.42 1.44 4
M/1 2.91 2.97 3.03 1.77 1.89 1.99 3/4
M/2 1.76 1.81 1.88 1.81 1.86 1.92 3
M/3 1.23 1.34 1.43 1.35 1.45 1.52 5
389
Table 4. Upper Ml/Upper M3 ratios for different species of Protatera from Eurasia.
(1) after Sen (1983). (Z) after Jaeger (1977).
Casablanca-M Z.63
Pul-e Charki (1) Z.70
Amama-Z (Z) Z.33
Ses Fontanelles Z.74
M/Z: This tooth follows the same pattern as the lower M1. However, a longitu-
dinal spur is present in most of the specimens.
M/3: This tooth displays an extremely simplified pattern, without distinguish-

able cusps or ridges.
Discussion:
The upper molars of P. almenarensis seem rather close to those of P. kabulense

from Pul-e Charki, AfghaniStan (Sen, op. cit.). They share a triangular shaped antero-
cone and a strongly reduced M3. Both are very different from E_. algeriensis from the
type-locality of the genus Amama-Z. However, the lower molars of P. almenarensis
are very different from those of E_. kabulense, with elongated M/1 diSplaying a very
broad anteroconid which resembles that of the extant Tatera indica. Nevertheless, a
relationship between the Iberian species and the last species seems not to be very
probable (the third molars of Tatera are not so much reduced as in P. almenarensis and
the upper molars still retain the posteroloph). -
A close comparison with other finds of Protatera in Spain is truly difficult, given
the scarce representation of this genus. Besides the Iberian finds, Protatera has been
also recorded in Ibiza, in the Balearic islands (Moya-Sola et al., 1984). The Ibiza
Protatera already displays a size larger than that of the continent, although the
M1/M3 relation is roughly similar. Protatera probably settled on this island during the
Messinian salinity crisis.
Besides Protatera, a second gerbil, Pseudomeriones, has been recorded in the

Casablanca-M fissure filling. This is a surprising find, since this genus had never been
recorded in western Europe. Curiously, the scarce material of Pseudomeriones from
Almenara exactly fits within the size and morphology of the type-species E_.
abbreviatus Teilhard from King-yan-fou (Kansu, China). On the other hand, the
Casablanca-M form is very different from the eastern Mediterranean species E_.
rhodh.ts from Maritsa (de Bruijn et al., 1970) and E_. tchaltaensis from Calta, Turkey
(Sen, 1977), which were already very specialized. While the find of Protatera could
maintain the doubts on the Asiatic or African character of the Casablanca-M fauna,
the presence of Pseudomeriones abbreviatus in this locality strongly suggest an Asiatic
origin.
A third supposed gerbil species, Epimeriones aff. austriacus Daxner-liock is also

present in the late Turolian of Can Vilella, in the Cerdanya basin. Epimeriones has a
completely different biogeographic character, its presence being restricted to central
and eastern Europe. This is the case, for instance, of the type-species E. austriacus
(early Turolian of Eichk'6gel), ~· progressus Kowalski from Podlesize (Poland), and the
poorly known E. dacicus Terzea from the early Pleistocene of Betfia-Xm.
Epimeriones aff-;- austriacus from Can Vilella seems very close to the Eichkogel
species, although it shows some advanced characters such as a higher hypsodonty and a
more simplified dental pattern. Therefore, two very different kinds of gerbil-like
forms settled the Iberian Peninsula in the late Turolian, one of central or eastern
European character (Epimeriones) and the other of African or Asiatic character
(Protatera).
390
'Muridae
The first murids in Europe (Progonomys cathalai) appear in a very low proportion
in some localities of the Cricetulodon zone (early Vallesian of Pedregueras 2C and Can
Llobateres). However, they are one of the commonest elements in the next biozone,
corresponding to the late Vallesian. In the Spanish basins, Progonomys appears split
into two different species, a cosmopolitan one (P. cathalai) and an Iberic variant ~·
hispanicus). This splitting coincides with a significant drop in the diversity of other
families (Cricetidae and Gliridae). In the late Vallesian of Terrassa, the P. cathalai
lineage is represented by large sized specimens not having attained the t6-=t"9 connec-
tion. A derivation of the early Turolian Parapodemus lugdunensis seems doubtful, the
latter species probably being an eastern immigrant.
The early and middle Turolian murid assemblages from the eastern Spanish
basins do not differ in a strong way from those in central Spain: Parapodemus
lugdenensis/~. barbarae, Occitanomys sondaari/0. adroveri, Valerymys vireti/V.
turoliensis. Nevertheless, in the early Turolian of Piera (Valles-Penedes) and La Celia
(Murcia) a second large sized Occitanomys, besides 0. sondaari, is present. The late
Turolian murid communities are characterized by their high diversity, with the entry
of Apodemus, Stephanomys, and Paraethomys. Curiously, a fourth form, Castillomys,
is already present in the fissure fillings of Salobrena and Casablanca-M, although it
remains absent in the fluvio-lacustrine isochronous sites such as Crevillente-6, Venta
del Moro, La Alberca, etc. until the early Pliocene. This faunal composition can be
extended to southern France (the "Ibero-Qccitanian province" of Hartenberger et al.,
1967) and even northern Africa (Stephanomys and Castillomys, see Coiffait-Martin et
al., 1985). Both Stephanomys and Castillomys are taxa derived from Occitanomys by
increasing stephanodonty (in the first case) and developing a longitudinal spur (in both
cases). The sudden appearance of Castillomys in the fauna of Casablanca-M and its
Mediterranean distribution suggest an eastern European or western Asiatic origin. On
the other hand, the distribution and first appearance of Stephanomys (§_. stadii from
Leberon) supports the idea of a western origin for this genus. In fact, some middle
Turolian populations of Occitanomys already show the tendency toward increasing
stephanodonty and the development of a longitudinal spur in the lower molars, a
condition fully attained in the late Turolian Stephanomys ramblensis.
In the case of Paraethomys, the most primitive members of this genus seem
related to the Asiatic species Karnimata darwini Jacobs (see Thomas et al., 1982). Its
presence in the two sides of the Mediterranean would indicate an isochronous settle-
ment of the two shores from the east. Moreover, some isolated teeth of a form close
to Karnimata have been recovered in the lowermost levels of the Molina de Segura
section (Fortuna), associated with Stephanomys ramblensis and Paraethomys
miocaenicus.
The transition to the Pliocene did not involve any special change among the
murid associations, although some cladogenetic processes affected the genera
Paraethomys (Paraethomys jaegeri from Gorafe-2, La Juliana, and Vilafant) and
Apodemus. In the latter case, an endemic lineage of large hypsodont populations
starts with Apodemus gorafensis from the Guadix-Baza basin. In the last member of
this lineage, A. agustii Martin-Suarez, the increasing hypsodonty caused some peculiar
features, such as the loss of the t6-t9 connection. While A. gorafensis started as a
vicariant reply of A. jeanteti from southern France, A. agustii shows some convergent
characters with the Rhagapodemus group (hypsodonty, thick enamel, etc.).
THE RODENT SUCCESSION
Late Ramblian ([bericus Zone)
The very early Aragonian (or late Ramblian, according to Daams et al., 1987) is
represented in the Valles-Penedes basin by the localities of Molt Calopa,
Costablanca-2, and Sant Andreu de la Barca (all belonging to the Ibericus zone). The
391
Table 5. Distribution of the Cricetid, Eomyid, and Glirid species in the late Ramblian
localities of the !hero-Levant subprovince (Valles-Penedes) and other
regions. + indicates presence, while (+) indicates inferred presence. Abbre-
viations: MC: Moll Calopa; CBl: Costablanca; SA: Sant Andreu de la
Barca. Other abbreviations as in figure z.
MC CB1 SA IC WE CE EE As Af
+ + + Pseudodryomys aff. ibericus + + (+)

+ Pseudodryomys aff. robustus +
+ Peridyromys murinus + + +
+ + Ligerimys elli:2ticus +
+ + Melissiodon dominans + + +
8
- - >75'/,
75'1.> > 50'1.
50'1.> > 25'/,
Af As
____ <2'1/o
Fig. 5. Zoogeographic congruence of the

late Ramblian rodent faunas from
the !hero-Levant subprovince with
the other recognized biogeographic
units. The different kinds of lines
indicate percentage of taxa shared
in common with a given bioprov-
ince. The encircled subprovinces
share more than 50% of their taxa
in common with the !hero-Levant
subprovince. Abbreviations are as
in figure z.
rodent associations of this phase are very poorly diversified, no more than five or six
species, lacking the cricetids of Oligocene origin (Eucricetodon, see table 5). The
dominant elements are the glirids of the subfamily Myomiminae (Peridyromys murinus
and Pseudodryomys ibericus). The sole cricetid is Melissiodon, while the eomyids are
represented by Ligerimys elli:2ticus. The sites in the Valles-Penedes basin share more
than the 50% of taxa in common with the !hero-Central basins, indicating that in this
time the Iberian Peninsula was a rather homogeneous biogeographic unit. The sole
difference with the central basins is the absence in the Valles-Penedes basin of the
endemic Myomiminae Armantomys, a taxon never found in this basin. However, there
is also a high degree of congruence with other European faunas such as Winthershoft-
West and Schnaitheim in Germany, therefore indicating a poor biogeographic differen-
tiation.
Early Aragouian (Primitivus Zone)
In this zone a very important increase of diversity takes place, with the entry of
the first CricetiDae (Democricetodon) and Cricetodontinae (Megacricetodon) in
Spain. Both genera have clear Asiatic connotations and their origin should be found
near some ancestral S:2anocricetodon stock. Together with Megacricetodon and
Democricetodon, a third cricetid, Eumyarion, is also present in the early Aragonian
beds of Can Mart1 Vell I and n (Vall~s-Penedes, see Agu!$tf, 1981) and Buiiol (Valencia,
see Daams and Freudenthal, 1974). On the contrary, Eumyarion is absent. in the early
Aragonian faunas of the Calatayud-Daroca basin.
392
Table 6. Distribution of the Cricetid, Eomyid, and Glirid species in the early
Aragonian localities of the Ibero-Levant subprovince (Valles-Penedes and
Araya basins) and other zoogeographic regions. Abbreviations: SM: Sant
Mamet; EC: El Canyet; CM-1: Can Marti Vell-I; CM-II: Can Marti Vell-II;
Ar: Araya. Other abbreviations as in figure z.
SM EC CM-1 CM-II Ar IC WE CE EE As Af
+ + + + + Democricetodo n aff. + (+)

hisEanicus
+ + + Megacricetodon minor + + (+)
Erimitivus
+ + Eumyarion weinfiirteri + + + (+)
+ + Ligerimys elliEticus +
+ + Ligerimys florancei + +
+ + PeridY!omys murinus + + +
+ + Pseudodryomys + + +
simElicidens
+ + MicrodY!omys sp. + + + +
+ + Glirudinus undosus + +
+ Glirudinus modestus + + +
+ Bransatoglis astaracensis + +

Ibero-Levant bioprovince with
other subprovinces during the early
Aragonian. For abbreviations see
figure z.
However, the dominant elements in these assemblages are the eomyids, repre-
sented in the Ibero-Levant province by two lineages, one of Iberian origin (Li~erimys
.elliEticus) and the other of European affinities (!:,. florancei). Among the glir1ds, the
Myomiminae are still dominant (PeridY!omys murinus and Pseudodryomys simElicidens)
but the Glirinae (Glirudinus) and the Dryomyinae (MicrodY!omys and Bransatoglis)
reappear. As in the case of the eomyids, Glirudinus is represented by two species, G.
gracilis and G. undosus. While G. gracilis is present in the Calatayud-Daro ca basin.
G. undosus is absent in the early Aragonian of central Spain. Another difference with
the Ibero-Central basins is the scarcity of MicrodY!omys in the Valles-Penedes basin.
Moreover, although a certain number of Iberian endemic forms are present in the
Ibero-Levant faunas of this age, there is a higher degree of congruence with the
western and central European faunas of similar age (for instance, Petersburch-Z, see
Ziegler and Fahlbusch, 1987). In fact, the three areas seem to form at this time a
rather homogeneous biogeographic unit (see figure 5). On the other hand, there is
little co~cidence, at the specific level, with the eastern European faunas, such as
Aliveri in Greece (see Benda and de Bruijn, 198Z).
393
Late Aragonian
The late Aragonian rodent faunas from eastern Spain display a degree of diver-
sity similar to that of the early Aragonian. The dominant elements are the
Cricetodontinae (more than 60%), with the genera Megacricetodon, Cricetodon, and
Hispanomys. The Cricetinae are very rarely represented. In this context, the locality
of Castell de Barbera is an exception, since Cricetinae and Cricetodontinae have
roughly similar percentages (probably this locality represents a very wet biotope
within a drier context; this interpretation is congruent with the abundance of the
Petauristinae in this site). Among the glirids, the Glirinae are highly diversified
(Glirudinus, Eomuscardinus, Muscardinus, and Myoglis), as well as the Dryomyinae
(Paraglirulus, Bransatoglis). The eomyids reappear, represented by Keramidomys. The
comparison of these late Aragonian faunas with those in central Spain (Escobosa, see
Sese, op. cit.) and western Europe (La Grive) indicates an equivalent degree of similar-
ity, although this biozone records the higher number of endemic species in this area
(Hispanomys dispectus, Cricetodon lavocati, Democricetodon ~· nemoralis). This
moment also records the highest number of Iberian endemics in the Valles-Penedes
basin (Megacricetodon ibericus, Fahlbuschia crusafonti, Muscardinus hispanicus). On
the contrary, the number of elements shared with central and eastern Europe is lower
Table 7. Distribution of the Cricetid, Eomyid, and Glirid species in the late
Aragonian localities of the lbero-Levant subprovince (Valies-Penedes basin)
and other zoogeographic regions. Abbreviations: SQ: Sant Quirze;
CB: Castell de Barber~; HI: Hostalets de Pierola (lower levels). The early
Vallesian site of Hostalets de Pierola (upper levels, HS) is also included
since it belongs to the Fahlbuschia crusafonti zone. Other abbreviations are
as in figure z.
SQ CB HI HS IC WE CE EE As Af
+ + + Cricetodon lavocati
+ + Hispanomys dispectus
+ + + + Fahlbuschia crusafonti +
+ + + Megacricetodon 1bericus +
+ + Megacricetodon minor minor + + (+)
+ + Eumyarion cf. medium + +
+ Democricetodon brevis nemoralis
+ + Anomalomys gaudryi + +
+ Cricetulodon hartenbergeri + (+)
+ + Muscardinus hispanicus +
+ + Myoglis meini + + +
+ + Paraglirulus werenfelsi + +
+ Keramidomys carpathicus + + +
CE
--- EE
At As
Fig. 7. Zoogeogrpahic congruence of the

lbero-Levant bioprovince with
other subprovinces during the late
Aragonian. For abbreviations see
figure z.
394
than in other biozones (Eumaarion medium! Myo~lis meiniC Keramidomys carpathicusci
etc.). Nevertheless, the 1fferences w1th t e Ibero- entrai provmce pers1ste
(absence of the endemic Myomiminae Armantomys and Tempestia and presence of
Eumyarion, Anomalomys, and Keramidomys). Finally, the eastern influence during
this time seems to have been rather restricted, since the Asiatic elements are absent
and there is a very low congruence with the Greek and Turkish faunas (Cricetodon/
Hispanomys in Spain, Turkomys/Byzantinia in Greece and Turkey, among other differ-
ences).
Early Vallesian
In the Valles-Penedes, where the stratotype is placed, the early Vallesian can be
subdivided into two very different phases. The first one (upper levels of the Hostalets
de Pierola section) displays the same faunal content as the late Aragonian assem-
blages, although Hipparion is already present. It clearly belongs to the Fahlbuschia
crusafonti zone.
The second phase (Cricetulodon zone) shows a completely different aspect. The
Cricetinae are usually the most common element, represented by the Cricetulodon
lineage (C. hartenbergeri-C. sabadellensis). On the other hand, the Cricetodontinae
are poorly represented, including new stages within the genera Hispanomys (H. thaleri)
and Megacricetodon (M. minor debruijni). Among the glirids, the Glirinae are again
the dominant element, but the Myomiminae reappear in the localities of Hostalets de
Pierola and Can Ponsic (Miodyromys hamadryas).
From a biogeographic point of view, there is also a clear difference within the
localities of the former biozone. Thus, no Iberian endemism is shared with the central
Spanish basins and there is only one endemic form (Hispanomys thaleri). The compari-
son with other European localities is difficult, since the early Vallesian is poorly
represented outside the Iberian Peninsula. Nevertheless, an idea can be drawn by
Table 8. Distribution of the Cricetid, Murid, Eomyid, and Glirid species in the early
Vallesian localities of the Ibero-Levant subprovince (VaUes-Penedes and Seu
d'Urgell basins) and other zoogeographic regions. Abbreviatiosn: CP: Can
Ponsic; CL: Can Llobateres; CPet: Can Petit; B: Ballestar. Other abbrevia-
tions are as in figure z.
CP CPet B CL IC WE CE EE As Af
+ + + Cricetulodon hartenbergeri/ + (+)

sabadellensis
+ + Hispanomys thaleri
+ + Megacricetodon minor + (+) +
debruijni
+ Eumyarion leemani (+) +
+ + Anomalomys gaillardi + +
+ 'Paraglirulus werenfelsi + +
+ Paraglirulus aff. lissiensis + +
+ + + Muscardinus hispanicus + + +
+ Eomuscardinus vallesiensis + + +
+ + Glis vallesiensis +
+ Bransatoglis astaracensis +
+ + + + Myoglis meini + +
+ Myomimus dehmi + + +
+ Eomyops catalaunicus (+) + + +
+ + Keramidomys C!!!l!athicus/ + +
pertesunatori
(+) Progonomys cathalai (+) (+)
395
IC EE

Ibero-Levant subprovince with
other bioprovinces during the early
Vallesian (Cricetulodon zone). For
abbreviations see figure z.
extrapolation from the latest Aragonian and late Vallesian/early Turolian sites of
France and Germany. In Germany, the early Vallesian is classically represented by the
localities of Howeneg and Eppelsheim, both having delivered only macromammals. On
the other hand, an interesting rodent association has been found in the locality of
Hammerschmiede (Jung and Mayr, 1980). Moreover, the locality of Dorn-Durkheim
displays an early Turolian association with micro- and macromammal remains that is
very close to the late Vallesian ones.
In a general way, there seems no strong differences between the supposed early
Vallesian faunas from the lbero-Levant province and those from western and central
Europe. The faunal composition of the Hammerschmiede locality (Jung and Mayr, op.
cit.) is especially significant in this way: six species of Cricetids (including the genera
Megacricetodon, Eumyarion, and Anomalomys), five species of glirids (with
Paraglirulus, Eomuscardinus, and Myoglis) and one eomyid species (Eomyops
catalaunicus). Thus, the early Vallesian faunas from eastern Spain display a higher
congruence with the western and central European ones than with those in central
Spain (Calatayud-Daroca and Duero basins), sharing most of the genera in the first
case and lacking the Iberian endemic species (for instance, Tempestia hartenbergeri)
of this age. Finally, it must be pointed out that at the end of this biozone the first
murids (Progonomys cathalai) appear in the site of Can Llobateres.
Late Vallesian
The transition from the early to the late Vallesian implied a very important
change among the rodent faunas. Therefore, the genera Megacricetodon and
Eumyarion disappeared, while Cricetulodon, Hispanomys, and Anomalomys persisted.
On the other hand, the change among the Glirid and Eomyid assemblages was even
more abrupt, with the exit of up to six different genera in the late Vallesian of the
Valles-Penedes basin: Bransatoglis, Myoglis, Eomuscardinus, Paraglirulus, Eomyops,
and Keramidomys. Thus, while the eomyids disappeared abruptly from the fossil
record of the Iberian Peninsula, the glirids were reduced to the few genera that are
now extant: Glis, Muscardinus, Myomimus, etc. On the other hand, the murids, repre-
sented by the genus Progonomys, became the dominant family among the rodents, a
situation that persisted until the early Pliocene.
As in the case of the early Vallesian, in the type area the late Vallesian can be
subdivided into two different assemblages. The first one, represented by the number
of localities in the Viladecavalls section and others in the Vall~s sector (such as Can
Casablanca& and Riu Ripoll) show a very poorly diversified rodent fauna, composed of
elements which survived the mid-Vallesian extinction: Cricetulodon sabadellensis,
Hispanomys thaleri, Anomalomys gaillardi, Myomimus dehmi. The sole novelty is the
dominance of the Iberian murid species Progonomys hispanicus, a form that appears
associated to the still-present E_. cathalai.
A second phase within the late Vallesian displays a somewhat different aspect
(Rotundomys bressanus zone). In this zone, the dominant forms are the Cricetinae of
396
Table 9. Distribution of the Cricetid, Murid, and Glirid species in the late Vallesian
localities of the Ibero-Levant subprovince (VaUes-Penedes basin) and other
zoogeographic regions. Abbreviations: Vc: Viladecavalls; TSA-n: Trinxera
Sud Autopista (Terrassa); TNA: Trinxera Nord Autopista (Terrassa); RR: Riu
Ripoll. Other abbreviations are as in figure z.
Vc TSA-n TNA RR IC WE CE EE As Af
+ + + + Hispanomys thaleri
+ Cricetulodon sabadellensis (+)
+ Anomalomys gaillardi + +
+ + Rotundomys bressanus (+) +
+ Myomimus dehmi (+) + +
+ + + + Progonomys aff. cathalai + + + (+)
+ + + Progonomys hispanicus +
WE .CE
IC ~IL(LEE
Af As

other bioprovinces during the late
Vallesian (Schizochoerus zone).
For abbreviations see figure z.

other zoogeographic bioprovinces
during the late Vallesian (bressanus
zone). For abbreviations see
figure z.
the genus Rotundomys (R. bressanus). In contrast with Cricetulodon, Rotundomys

presents a flat wear surface, a character also present in Microtocricetus. The remain-
ing elements of the latest Vallesian in the Valles-Penedes resemble those of the
previous biozone: Hispanomys thaleri, Anomalomys gaillardi. Nevertheless, the
degree of diversity of the very late Vallesian faunas from Terrassa is as low as those
i,n the Schizochoerus zone. From a biogeographic point of view, there are clear
differences with the central Spanish localities such as Masfa del Barbo, in the Teruel
basin, which yielded a very different rodent fauna; Rotundomys and Anomalomys are
lacking, while Kowalskia is already present and Hispanomys is represented by different
species (H. peralensis Vander Weerd). On the other hand, the rodent association from
Terrassa is very similar to that of the late Vallesian from western Europe, as it is
397
recorded in the localities of Soblay and the Amberieu section, in the Rhone valley
(Mein, 1984). Nevertheless, the late Vallesian from westem Europe shows some
differential characters compared with those in Spain. Thus, although the murids are
the dominant family, the persistence of the eomyids Eomyops and Keramidomys and
some glirid genera of middle Miocene origin (Myoglis, Paraglirulus) must be
mentioned. These data strongly suggest that the late Vallesian crisis had a latitudinal
character.
Early and MidcUe Tuolian
The early and middle Turolian is represented in the section of Piera, Camallera
(Emporda basin), Crevillente (Alicante, see de Bruijn et al., 1975), La Celia, and Casa
del Acero (the latter in the Fortuna basin, Murcia). In these five examples the faunas
show a rather homogeneous composition, very similar to that of the lbero-Central
basins. The degree of diversity is still very low, with three murids (Parapodemus,
Occitanomys, and Valerymys) and two cricetids (Hispanomys/Ruscinomys and
Kowalskia). This pattem strongly contrasts with the previous one in the late Vallesian
from the Valles-Penedes. Rotundomys and Anomalomys had disappeared and
Hispanomys is represented by a different lineage. The beginning of the Turolian
implied a strong unification between the whole Iberian faunas. The sole differential
element is Occitanomys n. sp., a large species of this genus that is present in the early
Turolian of Piera and La Celia. A second faunal difference is the persistence of
Hispanomys (H. adroveri) in the middle Turolian from Casa del Acero (Fortuna) and,
probably, from Crevillente-5, while in the Teruel basin the H. peralensis-H.
freudenthali lineage is followed by the very hypsodont Ruscinomys schaubi. Neverthe-
less, the general pattem shows a noticeable degree of homogeneity. During the
middle Turolian,. this homogeneity can be also extended to some western European
Table 10. Distribution of the Cricetid, Murid, and Glirid species in the early and
middle Turolian localities of the Ibero-Levant province (La Celia and
Fortuna basins) and other zoogeographic regions. Abbreviations: LC: La
Celia; CA: Casa del Acero (Fortuna). Other abbreviations are as in figure
z.
LC CA IC WE CE EE As Af
+ + Hispanomys sp./Hisl!anomY! adroveri

+ Kowalskia meini
+ + Parapodemus lugdunensisfP. barbarae + + + +
+ + OccitanomY! sondaari/0. adroveri + +
+ OccitanomY! n. sp. +
+ + Valerymys viretifY.. turoliensis + +
+ Eliomys truci + +
Af As

other bioprovinces during the early
and middle Turolian. For abbrevi-
ations see figure z.
398
localities, which display a similar composition (for instance, stade Cucuron, see Mein
and Michaux, 1979). This uniformity led Hartenberger et al. (1967) to recognize a
zoogeographic unit, the Ibero-Occitanian province, comprising the whole Iberian
Peninsula plus southern France. In fact, during the latest Vallesian and all the
Turolian there is an increasing biogeographic differentiation between western Europe
(Iberian Peninsula and France) and central and eastern Europe. This differentiation is
clearly reflected by the different murid composition in the two regions, and by the
persistence in central and eastern Europe of forms already extinguished in the western
localities. This is well recorded in the eomyids (Keramidomys as well as the early
Pleistocene genus Estramomys) and the glirids (persistence of Myomimus up to the
present while the western European basins are characterized by the presence of the
Dryomyinae Eliomys).
Late Turolian
As a result of the Messinian event, the late Turolian is a phase of large mammal
interchange in the entire Mediterranean basin (Moya-Sqla et al., 1984). Among the
rodents, this fact is reflected by an extraordinary increase in diversity, which espe-
cially affects the Murids and Cricetids. In the eastern Spanish basins, no typical
Iberian nor autochthonous endemism is present. On the other hand, although a higher
congruence with the rest of Spain and western Europe persisted, central European,
northern African, and Asiatic elements have been recorded also.
Western European elements: The late Turolian microfaunas from Spain are
mainly composed of species that are present in other parts of western Europe.
Elements of this kind are Apodemus gudrunae, Stephanomys ramblensis, Occitanomys
adroveri, Ruscinomys lasallei, Blancomys neglectus, and Eliomys truci. Some of these
forms have also been found in northern Africa. This is the case, for instance, of
Stephanomys, Ruscinomys, and Eliomys (localities of Ain-Guettara and Argoub
Kemellal, see Brandy and Jaeger, 1980 and Coiffait-Martin et al., 1985). A peculiar
case is that of Blancomys. This genus was first described in several Pliocene localities
from Spain (Arquillo-3, La Aldehuela, Villalba Alta, Sarri6n, Alcoy, Moreda, etc.) and
southern France (Sete and Mont-Helene, see Van der Weerd et al., 1977). Therefore,
Blancomys shows a typical western European distribution. Nevertheless, this form
seems not related to any of the Turolian cricetids present in the area. Recently,
Blancomys appeared also in the late Miocene of Casablanca-M. Since most of the
Casablanca-M elements show Asiatic affinities, we think also Blancomys to have had
an eastern origin.
Peri-Mediterranean taxa: In the late Turolian and early Ruscinian, two murid
genera, Paraethomys and Castillomys share a peculiar distribution around the
Mediterranean Sea. Therefore, Paraethomys is found in Spain, southern France,
Greece, Turkey, and northern Africa. Paraethomys miocaenicus was first considered
as an African immigrant in eastern Spain (Jaeger et al., 1975). However, an Asiastic
or at least eastern European origin has been assumed by other authors (for instance,
Aguilar et al., 1984).
EE
Af As

other zoogeographic bioprovinces
in the late Turolian. For
abbreviations see figure 2.
399
Table 11. Distribution of the Gerbillid, Cricetid, Murid, Zapodid, and Glirid species
in the late Turolian localities of the Ibero-Levant subprovince (Cerdanya
and Fortuna basins, Casablanca karstic complex) and other zoogeographic
regions. Abbreviations: CSB-M: Casablanca-M; MS: Molina de Segura
(Fortuna); CV: Can Vilella (Cerdanya). Other abbreviations are as in figure
z.
CSB-M MS-D MS-3 MS-8 MS-9 cv IC WE CE EE As Af
+ + + AEodemus + +
gudrunae
+ + + + + + Paraethom:rs + + + (+) +
miocaenicus
+ Karnimata sp. + (+)
+ + + + + SteEhanom:rs + + +
ramblensis
+ + Occitanom:rs cf. + +
adroveri
+ Castillom:rs + + + (+)
crusafonti
gracilis
+ + Cricetus kormosi + + + + (+)
+ Kowalskia sp. +
+ Ruscinom:rs + +
lasallei
+ Blancom:rs sp. + + (+)
+ Protatera (+) (+)
almenarensis
+ M:rocricetodon (+) +
cf. Earvus
+ Calom:rscus sp. + +
+ :Pseudomeriones (+) +
abbreviatus
+ EEimeriones aff. + +
austriacus
+ SminthozaEus + + +
janoss}'i
+ Muscardinus + + +
vireti
+ Eliom}'S truci + + +
A somewhat different case is that of Castillom:rs. C. crusafonti is a common

element in the rodent associations from southern Europe (it persisted until the early
middle Pleistocene of Cullar de Baza-C, in the Guadix-Baza basin). This genus is also
present in the localities of Maritsa, Greece (de Bruijn et al., op. cit.) and Calta,
Turkey (Sen, 1977) and in the site of Argoub Kemellal, in north Africa (Coiffait-Martin
et al., op. cit.). Moreover, C. crusafonti is already present in the late Turolian fissure
fillings of Casablanca-M (Agusti et al., op. cit.) and Salobreiia (Aguilar et al., op. cit.)
in eastern and southern Spain. The Casablanca-M rodent association is characterized
by its Asiatic affinities. Taking also into account the data from Maritsa and Calta, an
eastern origin for this taxon is proposed.
Central European elements: As stated, the presence of clear central European

taxa during the late Turolian has been recorded in the Pyrenees basin at La
Cerdanya. Therefore, the locality of Can Vilella yielded E¥imeriones aff. austriacus
and SminthozaEus janoss}'i, an association also observed in di ferent European sites.
400
African elements: Some of the elements present in Casablanca-M such as
Protatera and Myocricetodon are common elements of the north African micro-
faunas. Nevertheless, Protatera is also present in the late Miocene of Pul-e Charki,
Afghanistan (see Sen, op. cit.). Moreover, the Myocricetodontinae had a very wide
distribution during the Miocene, including western Asia (see de Bruijn et al., op. cit.).
Asiatic elements: The Asiatic character of the fauna from Casablanca-M is

fully confirmed by the presence of typical Asiatic elements such as Pseudomeriones
and Calomyscus.
The analysis of the late Turolian faunas from eastern Spain strongly suggests
that an important immigration event took place during the latest Turolian. This
immigration event was probably caused by the opening of new dispersal routes due to
the Messinian salinity crisis. However, the analysis revealed that most of these
immigrants were in fact of Asiatic origin and not of strictly African origin, as initially
proposed. In fact, it seems as if north Africa and central Asia were parts of the same
bioprovince. The Messinian regression probably allowed the extension of the limits of
this bioprovince northward, thus explaining the presence of Pseudomeriones,
Protatera, Myocricetodon, and others in Casablanca-M. A peculiar feature of this
aspect is the very different composition of the southeastern European faunas (Greece
and Turkey) which do not record any special change between the middle and late
Turolian faunas (see de Bruijn et al., op. cit.). This is a surprising datum, taking into
account the eastern character of some of the elements found in the Spanish late
Turolian. A way of explanation would be the possible different effects of the
Messinian crisis on both sides of the Mediterranean. In fact, the regression was
probably much more developed in the western basin than in the eastern one. The
eastern basin probably maintained a continous drainage from the fluvial system asso-
ciated with the Black Sea. On the other hand, the dessication effects in the western
Mediterranean probably were more intense, allowing the extension to the north of the
more steppe-like Afro-Asiatic bioprovince.
CONCLUSIONS
1) The Iberian Peninsula functioned very rarely as an homogeneous isolated

bigeographic unit. During most of the Neogene, the eastern basins of Spain displayed
a faunal content strongly influenced by western European taxa, lacking most of the
typically Iberian endemisms.
Z) The Pyrenees Range never acted as an effective barrier. In fact, during the
early Aragonian the whole European faunas (including western and central Europe) are
composed of the same basic taxa and can be assigned to a unique biogeographic unit.
3) In the Ibero-Levant province, the highest number of Iberian endemic and

autochthonous species is found in the late Aragonian. There is also a high degree of
congruence with faunas of western Europe, while there is a low similarity with faunas
of central Europe.
4) During the Vallesian, the Ibero-Levant province had the highest number of
taxa in common with the western and central European localities, while the degree of
congruence with the Ibero-Central basins was rather low. In the latest Vallesian, the
closest relationship is with the western European faunas.
5) The early/late Vallesian boundary marks an abrupt change among the rodent
associations, with the abrupt exit of many species that characterized the middle
Miocene.
6) The beginning of the Turolian suggests an unification of the microfaunas

from the Iberian Peninsula and southern France ("Ibero-Occitanian province"). More-
over, during the Turolian, a progressive differentiation between the western European
and the eastern-central European faunas took place; the western European faunas
being characterized by the persistence of taxa having a middle Miocene origin.
401
7) The entry of Asiatic taxa is restricted to the early Aragonian, the late
Vallesian, and the late Turolian. The entry of African forms is just restricted to the
late Turolian.
ACKNOWLEDGMENTS
This paper has been supported by the project "Bioeventos en el limite Mio-
Plioceno de las cuencas continentales del Levante espanol" (CICTY, PB86-058Z).
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Weerd, A. van der, Adrover, R., Mein, P., and Soria, D., 1977. A new genus and
species of Cricetidae (Mammalia, Rodentia) from the Pliocene of south-western
Europe. Proc. Kon. Akad. Wetens., Ser. B, v. 80(5), p. 429-439.
Wessels, W., Bruijn, H. de, Taseer Hussain, S., and Leinders, J., 1982. Fossil rodents
from the Chinji Formation, Banda Daud Shah, Kohat, Pakistan. Proc. Kon. Ned.
Akad. Wetensch., Ser. b, v. 83(3), p. 337-364.
Siisswasser-Molasse Niederbayerns. Zitteliana, v. 14, p. 3-80.
404
GUNDERSBEIM-FINDLING, A RUSCINIAN RODENT FAUNA
OF ASIAN AFFINlTIES FROM GERMANY
G. Storch
Forschungsinstitut Senckenberg
Senckenberg-Anlage 25
6000 Frankfurt/Main 1, Germany
0. Fejfar
Ustredni Ustav Geologicky

Malostranske N amesti 19
11821 Praha 1, Czechoslovakia
The Upper Ruscinian (MN 15) has a meager record in northern central Europe.
Our knowledge of Ruscinian rodents from Germany is restricted to the local faunas of
Wolfersheim near Frankfurt and- supposedly- of Gundersheim near Worms. They
are of some common interest because of their geographical position. Within Europe
they show affinities to well documented faunas both from eastern and southwestern
Europe, and because of their northern position they may as well exhibit affinities to
Russian and Siberian localities of similar latitudes which became known quite
recently. Thus, these local faunas may .contribute to biostratigraphical correlations
over these areas.
Micromammals from karst fissure fillings near Gundersheim, Rheinhessen, FRG,

have been first described by Heller (1936). Since that time, the stratigraphical
position of the fauna has been a matter of dispute. Heller originally placed it in the
Lower Cromerian as part of the Upper Pliocene and he compared it to localities like
Villany, Csarnota, and Beremend. Kretzoi (1956) considered the whole faunas as
Villanyian, and later (Kretzoi, 1962) regarded it as a composite of two clearly distinct
members; one belonging to the Upper Ruscinian (his Csarnotan) and being represented
by Glirulus pusillus (pl. 2, figs. ~6), Baranomys loczyi, Germanomys weileri,
Ungaromys nanus, and Mimomys hassiacus, and the other one belonging to the Upper
Villanyian represented by all remaining rodent taxa. This view has been widely
accepted, although not always on firm ground. H. Tobien, who has explored the
Gundersheim fissure fillings since the fifties, announced (Tobien, 1980) for the first
time the occurrence of the genus Trilophomys from Gundersheim. Obviously, this
indicates a previously unknown stratigraphical level.
We report here on a new rich and diversified Upper Ruscinian rodent assemblage,
Gundersheim-Findling. Along with W'olfersheim it represents an MN 15 fauna in
Germany and the more northern parts of central Europe on the whole. We have
reviewed the cricetids of Heller's original samples to demonstrate that they do not
include an Upper Ruscinian component, as was previously suggested.

Table 1. The Ruscinian rodent fauna from Gundersheim-Findling (MN 15), FRG.
Muridae
Rhagapodemus cf. ballesioi MEIN & MICHAUX 1970
Rhagapodemus frequens KRETZOI 1959
Apodemus atavus HELLER 1936
Micromys ~raeminutus KRETZOI 1959
Anomalomyidae
Prospalax kretzoii JANOSSY 1972
Trilophomyidae
Trilophomys schaubi FEJFAR 1961
Cricetidae
Baranomyinae
Cricetidae, nov. gen. canterranensis (MICHAUX 1976)
Baranomys longidens (KOWALSKI 1960)
Prometheomyinae
Stachomys cf. trilobodon KOWALSKI 1960
Germanomys weileri HELLER 1936
Arvicolinae
Propliomys hungaricus (KORMOS 1934)
Mimomys (Cseria) gracilis (KRETZOI 1959)
M. (C.) stehlini KORMOS 1931
~· (Mimomys) occitanus THALER 1955
M. (M.) hassiacus HELLER 1936
Cricetinae
Kowalskia cf. polonica FAHLBUSCH 1969
Zapodidae
Sminthozapus janossyi SULIMSKI 1962
Gliridae
Glirulus pusillus (HELLER 1936)
Glis minor KOWALSKI 1956
- ----
Eliomys intermedius FRIANT 1953
Muscardinus pliocaenicus KOWALSKI 1963
Muscardinus n. sp.
Sciuridae
Pliopetaurista pliocaenica (DEP~RET 1897)
Blackia woelfersheimensis MEIN 1970
Tamias orlovi (SULIMSKI 1964)
Gen. et sp. indet. I
Gen. et sp. indet. II
Our revision results in these taxa: Mimomys hassiacus (pl. 1, figs. 1-Z) is the
only species of the subgenus Mimomys. All specimens including the holotype are
equipped with cement, which contradicts Heller's diagnosis. The configuration of the
Linea sinuosa agrees with an evolutionary level seen in the European MN 16a zone
(Fejfar and Heinrich, 198Z, 1987). M. hassiacus is more primitive than M. polonicus of
MN 16b and M. pliocaenicus of MN 17. The Mimomys subgenus Cseria is represented
406
by four species: C. reidi, C. tornensis (pl. 1, fig. 4), and C. pitymyoides (pl. 1, fig. 7)
are characteristic of MN 17; the more primitive C. stehlini (pl. 1, fig. 3), according to
European standards, represents MN 16.
All lagurine specimens belong to Borsodia hungarica (pl. 1, fig. 6). The antero-
conid of an adult individual has a simple Arvicola-like shape which compares well with
material from European MN 17 localities. ID Germanomys weileri (pl. 1, fig. 5) we
include also Heller's supposed Ungaromys specimens. Germanomys is known from a
rather long time span, extending to the end of MN 16.
Thus, the revised Gundersheim fauna chronologically offers two interpreta-

tions. It may represent a lower MN 17 fauna with persisting primitive taxa, Mimomys
hassiacus, M. stehlini, and Germanomys weileri. Or else, it would represent two
chronological levels, MN 16a and MN 17. Contrary to previous suggestions, typical
taxa of European Ruscinian age (MN 14-15) are lacking.
The newly discovered Gundersheim-Findling fauna comprises Z7 rodent species

of 7 families (table 1). In contrast to Heller's material, typical cricetids of the Upper
Ruscinian are included: Trilophomys schaubi, Cricetidae (new genus) canterranensis,
Baranomys longidens, Stachomys cf. trilobodon, Mimomys occitanus, and Mimomys
gracilis. The occurrence of Mimomys hassiacus and M. stehlini, however, does not
seem to fit such an association.
Trilophomys schaubi (pl. 1, figs. 8-10) compares well with topotypical samples
from Ivanovce, and the rich material from Wt;ze (Kowalski, 1960; Sulimski, 1964)
surely belongs to that species too. The genus Trilophomys is rather common in MN 14
and MN 15 faunas of southwest to east-central Europe but so far is unknown outside
that area.
The species canterranensis (pl. 1, figs. 11-13) originally had been assigned to
Trilophomys (Michaux, 1976). Our Gundersheim-Findling material corresponds with
specimens from the type locality (MN 14) in the Rousillon basin, and the same species
is also known from Podlesice (MN 14), from W~ze and Ivanovce (MN 15). This taxon is
related to a group of small lophodont "arvicoloid" cricetids like Baranomys,
Microtodon, Anatolomys, and Celadensia which evolved during the Eurasian Turolian
and became widespread. None of these genera survived into the MN 17 zone.
Stachomys trilobodon (pl. 1, figs. 14-15) in the occlusal pattern, especially of its
first lower molar, resembles the genus Germanomys. Both occur in Gundersheim-
Findling and a comparison reveals, nevertheless, numerous differences in detail. The
folds alternate regularly and are U-shaped rather than V-shaped as in Germanomys,
the anterior labial and lingual folds of the last upper molar meet each other and are
not separate as in Germanomys, etc. Stachomys is known in Europe from its type
locality, W~ze, and neighboring Ivanovce (= "Leukaristomys vagui," cf. Fejfar 1961),
both Upper Ruscinian. Two further records are from south and west Siberia: Olchon
Peninsula of Lake Baykal where Stachomys is associated with Lophocricetus,
Microtoscoptes, Microtodon, Baranomys, and Dolomys (Mac et al., 198Z). This fauna
most likely is an equivalent of the European Lower Ruscinian, MN 14. The other
Siberian locality is Simbugino in Bashkiria where Stachomys occurs together with four
Mimomys species, Borsodia, and Plioctomys (J achimovich et al., 1977). This associa-
tion suggests an Upper Ruscinian age. These records favor the idea about a northeast
Asian origin of European "arvicoloid" cricetids (Fejfar and Heinrich, 1983). The
morphologically similar genus Germanomys occurs rather frequently in Ruscinian
faunas of east-central Europe such as W~ze, Ivanovce, Rebielice, and Osztramos-7. It
is missing in more southern and western localities, with Gundersheim and W'olfersheim
the only known occurrences in central and western Europe.
Propliomys hungaricus (pl. 1, figs. ZO-Zl) compares well with rich material from
the type locality Csarn6ta-Z (MN 15). Additional localities include W~ze and, repre-
sented by a single specimen, Mas Ge'n~gals in the Roussillon basin (Comet et al.,
1976). The closely related Propliomys ucrainicus is widely distributed in southern
407
1-21:~
a: 1 2mm,
~13
II 19 20 21
Plate 1. Cricetid and Anomalomyid molars from

Gundersheim near Worms, FRG, in occlusal
and labial (= fig. a) view. Specimens from
Heller's (1936) original sample (1-7), and
from Gundersheim-Findling (8-ZZ). 1,
Mimomys hassiacus, M/1; Z, M. hassiacus,
M/1-Z; 3, M. stehlini, M/1-3; 4, M.
tornensis, M/1; 5, Germanomys weileri,
M/1- Z, holotype; 6, Borsodia hungarica,
M/1-Z; 7, Mimomys pitymyoides, M/1-Z; 8,
Trilophomys schaubi, M/1; 9, T. schaubi,
M/3; 10, .I• schaubi, M3/; 11, Cricetidae, n.
g. canterranensis, M/1; 1Z, Cr., n. g.
canterranensis, M/Z; 13, Cr., n. g.
canterranensis, M/3; 14, Stachomys cf.
trilobodon, M/1; 15, .§_. cf. trilobodon, M3/;
16, Mimomys occitanus, M/1; 17, M.
gracilis, M/1; 18, M. hassiacus, M/1; 19, M.
stehlini, M/1; ZO- Z1, Propliomys
hungaricus, M/1; ZZ, Prospalax kretzoii,
M1-Z/ (ZZ on a reduced scale).
408
Russian equivalents of MN 15 faunas (Alexandrova, 1976) that again suggests an
eastern origin of the genus.
Mimomys (Cseria) gracilis (pl. 1, fig. 17) agrees with the type specimens from
Csarnota-Z and w1th samples from W~ze and Ivanovce. Cement is lacking, the
occlusal dentine pattern is confluent, the enamel band is rather thick and not differ-
entiated, and the Linea sinuosa is only slightly undulating. Two forms of the subgenus
Cseria that are normally found in Ruscinian localities of southern Russia and south-
western Siberia are Mimomys gracilis akkulaevae and M. baschkirika Siberia (Suchov,
1970).
The first lower molars of Mimomys occitanus (pl. 1, fig. 16) from Gundersheim-
Findling average slightly less than in a topotypical sample from Sete (MN 15).
Morphologically they exhibit the same features: rather thin enamel walls that are not
differentiated, lack of cement, weak elevations of the Linea sinuosa, enamel islet on
the first lower molar persisting long during wear, etc.
Thus, all of the cricetids mentioned so far clearly indicate a late Ruscinian
age. Non-cricetids strongly support this view. They are species typical of European
MN 15 faunas. Eliomys intermedius (pl. Z, figs. 9-10) as at its type locality, Sete, has
a second upper molar with two centrolophs and a first lower molar with a posterior
extra ridge and a long centrolophid. Pliopetaurista pliocaenica (pl. z, fig. 17)
compares well with specimens from Wolf.ersheim (Mein, 1970). Tamias orlovi (pl. Z,
fig. 15), Micromys praeminutus (pl. Z, fig. 4), and Rhagapodemus frequens (pl. Z, fig. Z)
cannot be distinguished from topotypical MN 15 material. There are also lineages
from Gundersheim-Findling that had been previously unknown from MN 15 localities
which support the biostratigraphical assignment. For instance, Rhagapodemus cf.
ballesioi (pl. z, fig. 1) is more advanced than the type specimens from the Lower
Ruscinian (MN 14) of Hautimagne (Mein and Michaux, 1970). It is somewhat larger
with the cusps higher and more inclined than the Hautimagne specimens.
Finally, tooth size of Kowalskia cf. polonica (pl. z, fig. 14) and Glis minor (pl. z,
figs. 7-8) would place Gundersheim-Findling rather low within MN 15, given a polarity
trend from small to large size.
But there are two advanced Mimomys species, M. hassiacus and M. stehlini, that
apparently do not fit this stratigraphic assignment. G\mdersheim-Findling thus
comprises two pairs of Mimomys species -- M. (M.) hassiacus and M. (M.) occitanus of
subgenus Mimomys, plus M. (C.) stehlini and M. (C.) gracilis of subgenus Cseria - each
of which differs in its evolutionary level. Such an association is unknown so far from
any Ruscinian locality of western and central Europe.
Unlike Mimomys gracilis, M. stehlini (pl. 1, fig. 19) is equipped with cement, the
dentine areas of the occlusal surface are only slightly confluent, the enamel walls are
rather thin, the Mimomys ridge is rather weak, and the enamel islet is lost during wear
in the apical third of the crown. Mimomys hassiacus (pl. 1, fig. 18) differs in a similar
way from M. occitanus. The labial aspect exhibits decidedly higher crowns of M.
stehlini and M. hassiacus with much higher elevations of the Linea sinuosa as
compared to tne more mesodont M. gracilis and M. occitanus. Morphologically the M.
stehlini molars correspond to the type specimen frOm San Giusto near Empoli (Chaline,
1974) and specimens from other MN 16 faunas like Arondelli, Seynes, and Moreda. The
M. hassiacus teeth would represent the European MN 16a faunal interval. M. hassiacus
is clearly more advanced than M. occitanus of MN 15 faunas and more primitive than
polonicus of MN 16b faunas. -
Thus, according to European convention the Gundersheim-Findling locality would

be of late Ruscinian age with contaminations by two younger, MN 16 elements. This
view is essentially altered by close comparisons with rich faunas from southern Russia,
the central Russian plains, and southwest Siberia, which are equivalents of the terres-
trial European Pliocene, MN zones 14 to 17 (Agadzhanyan, 1976; Alexandrova, 1976;
Jachimovich et al., 1977; Mac et al., 198Z; Suchov, 1970; Zazhigin, 1980). They come
409
Plate 2.. Rodent teeth from Gundersheim-Findling near Worms, FRG, in
occlusal view. 1, Rhagapodemus cf. ballesioi, M1/; 2., R.
frequens, M1/; 3, Apodemus atavus, M/1; 4, Micromys
praeminutus, M1/; 5, Glirulus pusillus, P4/; 6, G. pusillus, M1/;
7, Glis minor, P4/; 8, G. minor, M/1; 9, Eliomys intermedius,
MZ/; 10, ~· intermedius, M/1; 11, Muscardinus pliocaenicus,
M1/; 12, Muscardinus n. sp., M1/; 13, Sminthozapus janossyi,
M/1; 14, Kowalskia cf. polonica, M/1; 15, Tamias orlovi,
M/1,2; 16, Blackia woelfersheimensis, M/1,2; 17,
Pliopetaurista pliocaenica, M1,2/ (17 on a reduced scale).
from stratified fluviolimnic sediments, in part even from river terraces. These
assemblages are characterized by the coexistence of both primitive and advanced
Mimomys species. Of special interest are the supposed Ruscinian localities. They
extend from the Kishinev-Qdessa region (Kotlovina, Kryzhanovka, Morosovka,
Dolinskoe, Novaya Etulia, Livenzovka) through the middle Don (Uryv-1) to Bashkiria
and Kasakhstan east of the southern Ural Mountains (Akkulaevo, Simbugino, Beteke).
Primitive taxa which lack cement - Mimomys gracilis, M. baschkirika - are associ-
410
ated with advanced forms of two size categories which possess cement. In the Russian
literature the smaller advanced forms are assigned to M. stehlini, M. minor, and M.
livenzovicus. M. minor, and M. livenzovicus seem to be closely related to M.
stehlini. The larger forms in the literature are referred to M. coelodus, M. polonicus,
and M. pliocaenicus. Actually, they compare well with M. hassiacus and we suggest at
least close relationships.
To summarize, the rodents of Gundersheim-Findling represent a late Ruscinian

association of eastern European and Siberian affinities, which is so far unknown in
central and western Europe. These affinities are expressed by the occurrence of
species like Stachomys cf. trilobodon and in particular by the coexistence of Mimomys
species of different evolutionary grades. Since there are multilateral paleozoogeo-
graphical relations, this fauna can contribute to biostratigraphic correlations of
southwest Siberian with European localities, and above all to a reconsideration of the
correlation of Siberian faunas in terms of European MN units.
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the middle Don (russ.). Trudy Zool. Inst., v. 66, p. 58-97.
Alexandrova, L.P., 1976. Rodents of Anthropogene of the European part of the USSR
(russ.). Trans. Geol. Inst. Z91, 100 p.
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Mimomys occitanus-Mimomys stehlini et de Mimomys polonicus (Arvicolidae,
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Michaux, J ., 1976. D~couverte d'une faune de petits Mammif~res dans le Plioc"ene
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palaeont. Polon., v. 9(2), p. 149-244.
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412
DYNAMICS OF OLD WORLD BIOGEOGRAPmC REALMS DURING THE
NEOGENE: IMPLICATIONS FOR BIOSTRATIGRAPHY
Martin Pickford
Institut de Paleontologie
8, rue Buffon
75005, Paris, France
INTRODUCTION
The ultimate basis of biostratigraphy is organismal change, be it evolutionary or

distributional. In general the biostratigrapher is not directly concerned with what
caused the changes which he observes and uses for judging the deposition time of his
samples; he is more concerned with documenting those changes and understanding
their chronological connotations. The temporal scale that he constructs is valid and
useful without knowing what caused the changes on which it is based. However,
without an understanding of what lies behind these changes, biostratigraphy will
remain a descriptive pursuit. This paper is an attempt to get behind the biostrati-
graphic data and to understand what drives phylogenetic and distributional changes of
organisms. Such knowledge will not only provide a dynamic base, permitting us to
view biostratigraphic events within a framework of regional, global, and astronomical
change, but it may also allow us to use biostratigraphic data to reconstruct global and
astronomical events of the Neogene and Quaternary.
Among the various possibilities for changes in the biosphere, there are eight
which yield data of use to biostratigraphy (figure 1). These eight types of data fall
broadly into two categories, those using appearances in the fossil record and those
using disappearances. Of the two groups, the latter are inherently less reliable since
they are based on absence of evidence, which should never be thought of as evidence
of absence, but in many cases may provide evidence of a confirmatory nature.
Of these eight, three are dependent on changes in the distribution patterns of

organisms over time, four are based on autochthonous evolutionary events, and one on
extinction.
BIOSTRATIGRAPHY BASED ON AUTOCHTHONOUS EVOLUTION
In many cases autochthonous evolutionary events provide evidence of suffi-

ciently high quality that extremely refined biostratigraphic subdivisions can be made.
In tropical Africa, for example, tandem dental evolution among several lineages of
elephantids and suids occurred so rapidly that fine focus biostratigraphy is possible.
So fine is the resolution that suid biostratigraphy has become a recognized tool for
testing the validity of "absolute" dates based on paleomagnetic and radioisotopic
data. In every case in East Africa where biostratigraphy and geophysical dating have
produced incompatible results, the biostratigraphic data have always yielded the more
accurate initial age estimate.

Edited by E. H. Lindsay et a/.
D
w
physical/
2 3 4
A
2 3
B
Fig. 1. Eight ways that organisms in an area can
change to produce signals of value to biostra-
tigraphy. Top row: Four ways that organisms
can make an appearance in a local fossil record
(open circle in land mass (S)), from left to
right: 1. Immigration from a neighboring land
mass (P) across a physical barrier; z. Immigra-
tion from the same land mass, perhaps due to
ecological changes or to taxon pulse phenom-
ena; 3. Autochthonous phyletic speciation;
4. Autochthonous sympatric speciation. To the
left of each land mass (S) is depicted the
phylogenetic tree explaining each type of
appearance (black dot). Bottom row: Four
ways that cause the disappearance of a species
from an area (open circle in (S)), from left to
right: 1. Emigration without extinction;
z. Phyletic speciation; 3. Sympatric speciation
with extinction of parent species; 4. Extinc-
tion. To the left of each land mass (S) are the
phylogenetic trees which illustrate the dis-
appearances (black triangle) from the local
fossil record (open circle) in land mass (S).
However, a potential drawback to the use of autochthonous evolution in biostra-

tigraphy is the fact that it can seldom take into account contemporary geographic
variation. Two contemporary sites may contain faunas of slightly different aspect due
to geographic variability, but may be diagnosed as representing different time periods
by a biostratigrapher, despite their contemporaneity.
It is also possible that two superposed sites in a sedimentary sequence may yield
two distinct faunas which could be perceived as being time successive in the evolu-
tionary sense, when in fact they might have been contemporary geographic varieties
of organisms whose distribution patterns merely changed. A further drawback to
414
employing evolutionary events as the basis for biostratigraphy is that, in subdividing
an evolving continuum, paleontologists are essentially making arbitrary decisions as to
where taxonomic boundaries lie, and, in doing so, sometimes ignore ranges of variabil-
ity and occasionally cut a normal distribution into two or more fragments. Despite
these inherent problems, biostratigraphic schemes based on such an approach are often
robust, especially if attempts are not pushed beyond the resolving power of the basic
data.
BIOSTRATIGRAPHY BASED ON BIOGEOGRAPIDC EVENTS
Three categories of biostratigraphic evidence are based on changes in distribu-

tion patterns of organisms. Immigration of totally new lineages into a region are
often considered to be important biostratigraphic events, and in Europe have histori-
cally been employed to define the bases of several land mammal ages, including the
continental Oligo-Miocene boundary (the Brachyodus event), the base of the Orleanian
land mammal age (the proboscidean datum), and the base of the upper Miocene (the
Hipparion datum). There are some drawbacks to this approach to biostratigraphy, the
main one being that the immigrant lineage may not have colonized entire regions
simultaneously, in which case its appearance in different areas would have been
diachronous.
Disappearance by emigration of entire lineages from a region, leaving their

former ranges devoid of successors sometimes occurred, as for example the case of
equids in the Americas, but it seldom happened catastrophically, except perhaps on a
local scale. In any case, from a paleontological perspective it is unwise to base far-
reaching inference on absences from the fossil record. On a more localized scale,
there were undoubtedly frequent fluctuations in distribution patterns of organisms due
to many causes, and these might give the impression to the paleontologist that an
immigration or an extinction event had occurred If such a thing happened, it is possi-
ble that chronological significance could be inappropriately accorded to the event.
Since three of the eight biostratigraphic data sources have biogeographic

components, it is clear that many biostratigraphic concepts differ from biogeography
only by possessing the added dimension of time. The discipline of biostratigraphy is,
therefore, extremely closely allied to paleobiogeography. They are both dependent on
the same fossil data base; a. contribution to one automatically becomes a contribution
to the other; an improvement in understanding of one leads to an improvement in
understanding of the other; evidence from one discipline can be used to test hypoth-
eses erected in the other.
Without the biogeographic components, biostratigraphy is merely a descriptive

occupation; valid as a chronological tool, but lacking a dynamic explanatory founda-
tion. If we can determine what drives changes in the biogeographic world, we can
better understand what caused many of the faunal changes upon which biostra-
tigraphers base their correlations.
Aim
The aim of this paper, therefore, is to examine biogeography, using the neonto-
logic and fossil records of Africa and Eurasia as examples, to determine whether we
can tie down some of the regional, global, and perhaps astronomical events which may
have caused redistribution of faunas on a massive scale during the Neogene. Such
events have frequently been accorded importance by biostratigraphers who may
frequently be unaware of the ultimate causes of what they observe in the stratigraphic
column. By doing so, it is hoped that dynamic models can be built which can explain
why and how organismal distribution patterns changed. In addition, by reasoning in
reverse, we may be able to utilize the fossil record as a data source for reconstructing
local, regional, global, and perhaps even astronomical history.
415
EXTANT OLD WORLD BIOGEOGRAPHY
Latitudinally dependent organismal distribution patterns are such a prevalent

feature of the biosphere that biogeography was one of the earliest subdisciplines to
emerge in biology following the upsurge in interest in the living world in the nine-
teenth century. It is perhaps not merely coincidental that the founder of modern
biogeography, Alfred Russel Wallace, was also instrumental, if not decisive, in pro-
posing the Theory of Evolution by Natural Selection.
Wallace (1876) was the first to propose the subdivision of the biosphere into
latitudinally arranged biogeographic realms. In the Old World, excluding Australasia,
there are three Biogeographic Realms. The northernmost of these has been named the
Palearctic, and comprises Europe and much of Asia north of 30°N. It also includes the
Maghreb, in northern Africa. To the south of the Palearctic occur the Ethiopian
Realm (Africa without the Maghreb) and the Oriental Realm (India and Southeast Asia)
(figure Z).
Wallace noted that the latitudinal arrangement of these realms was essentially a
widespread response by organisms to latitudinally expressed global climatic condi-
tions. The Ethiopian and Oriental Realms are essentially tropical to sub-tropical,
while the Palearctic is comprised of the temperate, boreal, and arctic belts. In terms
of organismal dynamics, the boundaries between the Palearctic Realm to the north
and the Ethiopian and Oriental Realms to the south coincide closely with the boundary
which separates climates with a winter-summer regime to the north from those with
wet season/dry season alternations to the south.
From the point of view of terrestrial mammals, it is important to note two

major consequences of this subdivision. Firstly, in the Palearctic Realm, breeding is in
phase with a winter-summer cycle, whereas in the Ethiopian and Oriental Realms it is
tied into wet season/dry season cycles, except in places where there is little season-
ality such as in tropical forests. Secondly, plant growth is markedly temperature
dependent in the Palearctic with the main growing period lasting for about three to six
months each year, with up to three months without significant growth (figure 5).
Fig. z. Modern Biogeographic Realms plotted on a world map

cut into 1 Z gyres. Biogeographic Realms are as
follows: A - Australasia; 0 - Oriental; E - Ethiopian;
P- Palearctic; N- Nearctic; T- Neotropical. The
boundaries between tropical realms and high latitude
ones coincide more or less with the zone in which a
winter-summer climatic cycle to the north gives way
to a wet season/dry season climatic cycle to the
south.
416
There are several strategies employed by Palearctic animals to overcome
seasonal fluctuations in fresh food supply, including hibernation, migration in response
to the annual cycle of temperature fluctuations, and adaptations to locate and digest
whatever food supplies remain during the winter season. In the Ethiopian and Oriental
Realms, in contrast, growth periods are not limited by temperature fluctuations, but
by rainfall patterns; consequently strategies to overcome seasonal shortfall in food
resources comprise estivation, migration in response to precipitation patterns, and
adaptations for obtaining and digesting underground food resources and other scarce or
durable items. However, in huge areas of the tropics fresh plant food is available all
the year round (figure 5).
These two points are important to understand, since they have implications for
the biostratigraphy of Europe. Consider African mammals adapted to a food supply
fluctuating in phase with a wet season/dry season cycle, and timing their reproduction
to the same cycle. If such species were to colonize the Palearctic Realm, they would
have to modify their annual feeding strategy as well as the timing of their reproduc-
tion to accord with a winter-summer cycle. Their response to the Palearctic environ-
ment would become temperature dependent rather than precipitation dependent.
While this has undoubtedly happened sometimes in the past, it probably was not very
common. Otherwise the modern Palearctic and Ethiopian/Oriental Zoogeographic
Realms would not be as distinct from each other as they are. One only has to think of
such mammals as the woolly mammoth and woolly rhinoceros to find examples of
former Palearctic inhabitants whose ancestral stocks probably had tropical origins.
Conversely, some deer species have been able to colonize the tropics, their fore-
runners presumably having originated in the Palearctic. Nevertheless, many lineages
of mammals have apparently been unable to make the transfer, which is why to this
day the zoogeographic realms are still populated by distinctive faunas. Interestingly
enough, meat-eating Carnivora, being dependent on food resources (animal proteins
and other products) which are essentially comparable in all biogeographic realms,
comprise one group of mammals which show little respect for zoogeographic bound-
aries. Herbivores, being directly dependent upon vegetation, are generally restricted
to a single zoogeographic realm or to realms which are similar in seasonal structure,
such as the Oriental/Ethiopian pair, and the Palearctic/Holarctic pair.
There are, therefore, powerful biogeographic constraints on the distribution of

organisms at the scale of the biogeographic realm. From the point of view of biostra-
tigraphy, there are many implications which must be considered. Firstly, biochrono-
logical correlations between neighboring biogeographic realms may be difficult or
impossible, because of the pervasive differences which exist between neighboring
realms. Consider trying to correlate fossiliferous strata deposited during the past ten
years in Sweden, Italy, and Kenya. Even though the deposits would have accumulated
during exactly the same period, there would be almost no biological similarity between
their fossil contents. A biostratigraphic correlation would be impossible.
Now consider the role of latitudinal biogeographic effects during the Miocene
and try to assess the potential for obtaining correlations between East Africa,
southern Europe, and northern Europe.
Secondly, it is likely that the various zoogeographic realms have existed for long
periods of time, even if the faunas and floras within them have evolved considerably.
The astronomical and global conditions which define the boundaries of the realms
within the biosphere have undoubtedly been in existence for at least the entire
Cenozoic if not longer. It is equally likely that the extent and distribution patterns of
these conditions on the surface of the globe have not remained static. Given that
global and astronomical factors have varied during the Cenozoic, there were probably
times when the proto-Palearctic zone was more restricted in latitudinal extent than it
is now, and there may have been periods when it was more widespread.
If this is so, then it is of particular importance for European biostratigraphers,

because their fossil samples are situated at the latitudes most likely to have experi-
enced successive fluctuations in biogeographic boundaries from Palearctic to
417
Ethiopian and back. At the equator, in contrast, it seems safe to assume that condi-
tions were always tropical, unless the position of the equator changed.
TAXONOMIC DIVERSITY GRADIENTS (WALLACE'S RULE)
Wallace was the first to observe that the taxonomic diversity of numerous
groups of organisms is greatest near the equator and drops as higher latitudes are
approached. A tally by latitude of the number of species in a family or higher rank of
organisms yields a curve described as a taxonomic diversity gradient. This latitudi-
nally generated diversity curve should not be confused with ecological diversity or
species richness in habitats or ecological regions. The concept of taxonomic diversity
gradients as employed by biogeographers is entirely independent of local and regional
ecological parameters, being solely a function of latitude, regardless of what habitats
occur at each latitude. It is necessary to keep the scale of the model in mind when
working with latitudinal taxonomic diversity, and not to confuse it with local or
regional taxonomic richness.
Wallace's Rule applies to so many groups of organisms that it is probably an

aspect of a fundamental relationship between life and energy. Many taxonomic
diversity envelopes are of similar shape to the curve which describes global solar
energy capture (Stehli, 1968) (figure 3). The biosphere appears to respond in two ways
to energy input. Firstly, biomass production is closely related to energy input; the
more energy that enters a biological system, the greater will be the biomass produc-
tion. Secondly, biological diversity seems to be directly related to energy supply; the
more energy available, the higher the biological diversity. Not only does this second
phenomenon apply to taxonomic diversity, it also seems to apply to variability within
populations of single species For example, polymorphisms, be they molecular or
karyological, seem to be more varied in the low than in the high latitudes.
20,...------------,
100
---·'\
"
'- >.
1/ ·"·
QJ
E'
~12 QJ
c:
.I \\
.....
0
QJ
'- Jlso
1: 4 0
E I •
\ "'
0~
~
90 0 90 90 0 90
N s N s
Fig. 3. Relationship between taxonomic diversity
gradient (left frame) and solar energy capture
(right frame). In both, the peaks of the
envelopes are over the equator, and the low
value ends of the limbs are at high latitudes.
Left-hand frame is data for mosquito genera
(Stehli, 1968) and right-hand frame is solar
energy capture as percentage of insolation
incoming to the globe (also Stehli, 1968). The
shape of the envelope for mosquitoes is
typical of numerous groups of organisms, be
they plants, invertebrates, or vertebrates, and
the pervasive presence of this type of curve
in the biosphere suggests that a close rela-
tionship exists between taxonomic diversity
and quantities and disposition of solar energy
input into the globe.
4·18
20
"'
X
....."' ,2
.....0
c...
QJ
.c
5 t.
z
90 0 90
N s
Fig. 4. Diagram to depict effects of fluctuations in
solar energy input into the globe. With
increases in solar energy receipts, the taxo-
nomic diversity envelope inflates (open
arrows) whereas with decreases in solar
energy capture, the taxonomic diversity
envelope deflates (black arrows). Note that
envelope inflation results in an increase in the
latitudinal range of the group of organisms
under consideration (open arrows at end of
envelope limbs). A fossil locality at or near
this latitude (70° in this hypothetical case)
would experience ingression and regression of
taxa as solar energy levels fluctuated. For
many groups of organisms, southern Europe
lies at a latitude near the ends of taxonomic
diversity envelopes. Thus, during the
Neogene it experienced many cases of faunal
immigration and emigration which may have
been due to such fluctuations in levels of
solar energy capture. Note that a deflation
of the envelope means loss of taxa, i.e., an
extinction event, whereas an inflation of the
envelope means an increase of taxa, i.e., a
radiation. Radiations and extinctions caused
by such events would be out of phase with
each other. At present, the Holocene global
warming may well be driving the active
radiation of the Cercopithecus group monkeys
and Cichlidae of tropical Africa.
If the global solar energy capture varied over geologic time and if Wallace's Rule
applied in the past as it does today, then the following deductions can be proposed. An
increase in solar energy capture would correspond with an increase in biomass produc-
tion and taxonomic diversity, while a decrease in solar energy capture would corre-
spond to a decrease in biomass production and taxonomic diversity.
In the high latitudes, where solar energy capture fluctuates widely during the
year (figure 5), biomass production and diversity of organisms increases and decreases
cyclically in phase with the seasons. Biomass production varies as a direct conse-
quence of fluctuations in insolation. It must be admitted, though, that annual
taxonomic fluctuations are primarily due to migrant visitors. The annual cycle is too
short for taxonomic diversity to increase and decrease by speciation; instead, myriad
mobile species of tropical and sub-tropical organisms have incorporated annual
migratory visits to high latitudes into their repertoire of life strategies. Many truly
Palearctic taxa merely enter a state of torpor, or suspended animation during the
winter months, and thereby withdraw temporarily from the active biosphere, to re-
419
Fig. 5. Biomass production levels at various latitudes. Note the
relatively stable levels near the equator which contrast
strongly with the seasonal fluctuations in the northern hemi-
sphere. The high latitudes in the south have a uniformly low
biomass production on land because there is little unglaciated
land area in the far south of the globe. Seasonal migrations of
mobile tropical and sub-tropical taxa to the high north lati-
tudes lead to annual inflation and deflation of the taxonomic
diversity envelope of the high latitude part of the biosphere.
This should not be confused with Wallace's Rule, although the
analogy is remarkable and the two may be related because of
their ultimate link to levels of solar energy capture. The
general shape of the north-south curve depicting winter
biomass production closely resembles many taxonomic divers-
ity gradients. (Data from Tucker et al., 1986; Malingreau et
al., 1987.)
emerge the following year as fully participating members of the taxonomic society.
This annual high latitude taxonomic diversity fluctuation is not, strictly speaking, an
aspect of Wallace's Rule, but there may be a connection between this phenomenon and
latitudinal diversity gradients, since both are ultimately dependent upon quantities of
solar energy capture.
In the tropics, however, solar energy capture fluctuates very much less during
the year (figure 5), so that taxonomic diversity is more stable, but if long term
increases or decreases in solar energy input were to occur, there would probably be
appropriate responses from the biosphere. In effect, if the solar energy gradient
envelope was inflated, taxonomic diversity gradients would follow suit (appropriate
biosphere response). Conversely, if the solar energy envelope deflated, so would
taxonomic diversity envelopes, which would be observed by a biologist as a decrease in
diversity (figure 4).
From biogeographic and biostratigraphic perspectives, the relationship between

solar energy capture and taxonomic diversity are important for several reasons.
Firstly, if solar energy capture were to increase from present levels, then the
limbs of the envelope which describes the gradient of solar energy capture would move
toward higher latitudes. This would mean that the higher latitudes would become
more tropical than they are now. If increases in solar energy were high enough, then
conditions in the Mediterranean region could well become similar to those in the
northern parts of the Ethiopian and Oriental Biogeographic Realms (i.e., it would
420
become sub-tropical rather than temperate), and southern Europe would witness an
influx of Ethiopian and Oriental organisms, access pathways permitting (figure 6). At
the same time a retreat northward of Palearctic denizens would occur. In view of the
immense climatic fluctuations which occurred during the Plio-Pleistocene, it is impor-
tant to keep this relationship between solar energy capture and taxonomic diversity in
mind.
Fig. 6. If a shift in the position of the equator

occurred, as postulated by Koeniguer (1987)
and Termier and Termier (1986), it would
result in a shift in the position of the taxo-
nomic diversity gradient. Evidence from
palms (top frame) and anthropoids (bottom
frame) as well as numerous other tropical
African organismal groups indicate that such
a repositioning of the tropical belt may have
occurred. It could be inferred that a shift in
orientation of the equator may have taken
place during the Miocene. If all else, such as
levels of solar energy capture, remained
stable, the postulated shift in equatorial
position and the tropical belt with it, taken on
their own, would result in the establishment
of Ethiopian conditions further north than
they exist now. If this happened, southern
Europe would be suitable for colonization by
African and Oriental faunas without their
having to undergo major adaptations. Note
that, if this model is correct, southern Africa
would become less tropical than it is now,
making it unsuitable for habitation by
primates and other strictly tropical
organisms. Note also that if the shift in the
eastern Hemisphere was to the north, in the
western Hemisphere the shift would be
southward (see figure 7).
421
Fig. 7. A southward shift in the position of the
tropical belt, and by inference the equator, in
the western hemisphere would result in the
southern tip of South America becoming more
tropical than it is now. This would make it
suitable for habitation by Platyrrhines. The
Santacrucian (early Miocene) of Patagonia
yields abundant Platyrrhines, a fact which
supports this model. For further explanation
read caption to figure 6 and the text.
Secondly, some of the Pleistocene extinctions may well be related to a shrinkage

of the taxonomic diversity envelope caused by reductions in levels of solar energy
capture, and, as such, they may have been worldwide in effect. Conversely, some of
the taxonomic radiations of the Plio-Pleistocene, including those of the suids,
cercopithecids, and bovids, may be related to extended periods during which solar
energy input increased above previous levels. The present-day proliferation of
Cercopithecus monkey species and cichlid fishes in tropical Africa may well be
examples of responses to such an increase in solar energy input since the Holocene.
Thirdly, because in numerous groups of organisms the peak of the taxonomic

diversity envelope is associated with the equator, the position of peaks of ancient
diversity gradients, if they can be determined, may be used to define the positions of
the equator during the past, as was realized by Stehli (1968) (figures 6 and 7).
ASTRONOMICAL ASPECTS
Changes in the amount and distribution of solar energy received by the Earth
could be due to a variety of causes. Fluctuations in solar energy output is an obvious
example. Change in the orbit of the Earth around the Sun would result in modified
energy receipt by the Earth. Even if solar output and orbital parameters remained
constant, there might still be changes in the distribution of solar energy on the surface
of the globe, which would have profound effects on the biosphere. For instance, if the
axis of rotation of the globe were inclined at a different angle to the ecliptic, there
would be major changes in the degree of seasonality on the globe. If it were oriented
at 90° to the ecliptic, there would be little or no seasonality, and the length of the day
would be constant over the entire globe. If the angle were greater than at present,
polar winters would be even more extreme than they are now. A further possibility is
that the axis of spin of the globe might change (Lavallard, 1988), even if the globe
retained its other relationships to the solar system. If this happened, the position of
the equator would change, and with it the distribution of the biogeographic realms on
the surface of the globe (figures 6 and 7).
Two items from the above list are testable by recourse to data from the fossil
record. These are fluctuations in solar energy output (figure 4) and changes in the
position of the pole of rotation (figures 6 and 7). Other factors, for the time being,
422
are assumed to have remained constant. Two aspects of biogeography are pertinent
here; the first is the pattern of distribution of organisms on the surface of the globe
(i.e., traditional biogeography) and the second is taxonomic diversity (Wallace's
Rule). Both of these are considered to be of potential for studies of fluctuations in
solar energy flux, and polar repositioning because both are ultimately controlled by
a) the amount of solar energy captured by the globe, and b) the distribution of this
energy to the biosphere.
o18o DATA FROM SEAFLOOR. CORES

Oxgyen 18 data obtained from benthic foraminifera of the Pacific and Atlantic
Oceans (Miller and Fairbanks, 1985) yields a curve which represents fluctuating values
of Oxygen 18 relative to Oxygen 16 throughout the Ce~oic. The ratio of Oxygen 18
to Oxygen 16 is temperature dependent, and thus the o 0 curve can be visualized as
a paleotemperature curve for the waters in which the foraminifera were growing. It
has alref~Y been suggested (Pickford, 1987c) that there are some correlations between
major 8 0 events during the Neogene and changes in continental faunas (figure 8).
s1Bo
3·0 2·0
cooler ------,-- Temperature - h o t t e r
..,~
Vl
_, -25
~ my ..,
Vl
;::)
z
0
N
~
:!S
.....
:E
Fig. 8. o18o curve of benthic foraminifera of the Pacific and Atlantic

Oceans. Superimposed on the curve are tropical African fossil
mammal localities. To the left is the biostratigraphic subdivision of
the East African Neogene, core faunas (=niveaux reperes) being
identified within the curve (Pickford, 1981), and on the right is the
column of Neogene mammal zones of Europe. (Oxygen 18 curve from
Miller and Fairbanks, 1985.)
423
It is not known what caused the temperature of ocean waters to change,
although it is sometimes assumed to have been due to changes in quantities of solar
energy capture or to rearrangements of circulation patterns of oceanic waters follow-
ing changes in the configuration of continental plates. The mid-Miocene cooling event
(Miller and Fairbanks, 1985), for example, is said to be related to the establishment of
the circum-Antarctic current and the subsequent growth of the Antarctic ice sheet.
If, on the other hand, climatic change was related to changing amounts of solar energy
capture, then taxonomic diversity envelopes would be expected to have inflated or
deflated according to rises and falls in paleotemperature. While some of the paleonto-
logical data suggest that for some of the events this might indeed have been the case,
it does not seem to satisfy all of the paleobiogeographic data, especially that concern-
ing the position of the peak of ancient diversity gradients.
PLIQ-PLEISTOCENE FLUCTUA110NS IN TAXONOMIC DIVERSITY
Inflation and deflation of taxonomic diversity envelopes is considered to result

from fluctuations in the quantity of solar energy captured by the globe as already
described above. In favorable circumstances, the fossil record is complete enough
(figures 9 and 10) for ancient diversity levels of well sampled fossil organisms to be
reconstructed for several key latitudes. Among African mammals which fall into this
category are the Suidae, Proboscidea, and Cercopithecidae (figure 10), but other
groups would be just as amenable to this kind of analysis. These three groups reveal a
common pattern during the Plio-Pleistocene. Firstly, the limbs of the diversity
envelopes reached further toward high latitudes than they do now, and the diversity
peak at the equator was greater during the Plio-Pleistocene than it is now. This
symmetrical inflation of the envelope during the late Pliocene and its subsequent
deflation is considered to be in accord .with the suggestion that the levels of solar
energy received by the globe have fluctuated during the Plio-Pleistocene while the
equator occupied more or less its present position.
Fig. 9. Map of the western part of the Old World

showing the fossiliferous areas which have
yielded the data used to compile the taxo-
nomic diversity gradients illustrated in figure
10.
424
t
10 SUI FORMES
_,,'' ~-- ...,. Omy
I , ~ •'' ' ' I , , , , I

90 0 90
" 8
.....~ PROBOSCIDEA
.....0
...01 4 ~2·6my
.c
:·-~--~Om~
E
:::1
:z /
0 ' I
90 0 90
N s
.....0
...01 8
.c
E
:2.
0
90
s
Fig. 10. Modern and fossil taxonomic diversity enve-
lopes for the late Pliocene/early Pleistocene
and Recent mammalian groups: Suiformes,
Proboscidea, and Cercopithecoidea. Note
that in all three cases the taxonomic diversity
2..6 Ma was greater and the latitudinal exten-
sion higher than they are now. Since the
changes were apparently symmetrical on
either side of the present-day equator, this
data suggests that 2..6 Ma the tropical belt
and the equator were more or less in their
present positions, and that the main causes of
the subsequent deflation was a lowering of
global solar energy capture, which resulted in
loss of taxa, i.e., extinction. Other organis-
mal groups show the same pattern of change,
despite the hazards of the fossilizing and
collecting processes, which are usually con-
sidered to result in taxonomic under-repre-
sentation in the fossil samples.
If inflation of the taxonomic diversity envelope occurred, its limbs would move
toward higher latitudes. From the point of view of the biostratigrapher working in the
Mediterranean Basin, this would be perceived as an increase in diversity of Ethiopian
taxa in the Mediterranean fossil record. Conversely, a deflation of the diversity
envelope would be perceived as a decrease in diversity of African mammals in the
southern European region. Ethiopian mammals such as Hippopotamus, Crocuta, and
Macaca, which colonized Europe during the Plio-Pleistocene only to become extinct
later on, illustrate the point.
425
MIOCENE BIOGEOGRAPHY
Different in nature from the inflation and deflation of taxonomic diversity

gradients is the situation whereby the position of the peak of taxonomic diversity
envelopes apparently changed (figures 6 and 7). During the Miocene, both limbs of the
diversity envelopes were apparently further to the north than they are now
(Koeniguer, 1988; Termier and Termier, 1987), suggesting that, in Africa, the peak was
about Z0° north of its present position. Evidence for this reconstruction comes from a
variety of different groups of organisms, including marine sponges, littoral mangroves,
terrestrial palms, and several groups of vertebrates, including fishes, turtles,
crocodiles, and mammals. This suggestion is supported by the distribution patterns of
various groups of organisms, including Primates and Giraffidae among others (figures
11 to 14).
From the perspective of the Neogene biostratigrapher working in Europe, a

northward shift in the diversity peak would be recorded as an increase in the diversity
of African taxa in the southern European sedimentary sequence. In fact, if the worker
confined himself to the study of the European sequence alone, it would be difficult, if
* Miocene palms
• Miocene mangroves
Fig. 11. Map of part of the Old World with the distribu-
tion of Miocene and Recent palms and man-
groves. Note the northward expansion of the
ranges of these essentially tropical and sub-
tropical plants. Mangroves in the middle
Miocene of Kurosedani, Japan, are about
1000 km north of the nearest living examples,
and their presence in Japan suggests a heating of
coastal waters by 10° in the summer and ZO in
the winter compared to modern sea water
temperatures (Karyu et al., 1984). In Europe,
mangroves grew on the north shore of the
Mediterranean Sea (Koeniguer, 1987) and palms
grew north of the Black Sea (Chepalyga, 1985).
The vast longitudinal extent of these changes in
distribution patterns (Portugal in the west to
Japan in the east) indicates that we should be
searching for a global scale cause, rather than a
parochial Mediterranean one.
426
ANTHROPOIDEA
Extant
Miocene •
Fig. lZ. Distribution of Miocene Anthropoidea. The

northward expansion of the range of anthropoids
is evident in this map, and accords well with the
model in which the Miocene tropical belt was
about Z0° north of its present position. Note
that if this model is correct, then South Africa
becomes inhospitable to anthropoids, whereas
southern South America becomes habitable to
them. These suggestions accord with currently
available fossil data. (N - postulated position of
Miocene North Pole; S- postulated position of
Miocene South Pole.)
GI RAFFIOAE
Extant c
Miocene •
Fig. 13. Distribution of Giraffidae during the Miocene

accords with the biogeographic boundaries
inferred from anthropoid distribution (see figure
lZ).
427
PLEURODIRA
Extant D
Miocene •
Fig. 14. Distribution of Miocene Pleurodiran turtles.

Note the northward expansion of their range in
Eurasia of a similar nature to that seen in
anthropoids, giraffids, mangroves, palms, and
other tropical to sub-tropical organisms.
Numerous other groups of tropical organisms
show the same pattern of distribution during the
Miocene. (Data from de Broin, 1987.)
not impossible, to determine whether the diversity changes were due to a shift in the
position of the diversity peak, or to an increase in global solar energy capture, or to
some other cause perhaps of a more parochial nature. This is why it is necessary to
examine the possibilities at various scales, ranging from local and regional to global.
In order to perceive global effects, it is necessary to obtain samples from various
latitudes on both sides of the equator (figures 9 and 10). Only then is it possible to
determine whether the changes were due to increases in solar energy (effects would be
symmetrical on either side of the equator) or to change in position of the tropical belt
(effects would be asymmetrical with respect to the present-day equator), or whether
the effects were the result of regional or local changes, rather than global ones.
For example, during the Plio-Pleistocene increases in the diversity of monkeys in

East Africa were contemporary with increases recorded in South Africa (30°S) and
North Africa (30°N) (figure 10) (Pickford, 1987a), from which it is suggested that, with
respect to the modern equator, a symmetrical inflation of the taxonomic diversity
envelope took place. In contrast to this example, the increases in diversity of
Anthropoidea in Europe during the Miocene was contemporary with a decrease in
diversity in East Africa (Pickford, 1986b) (figure 18). In South African deposits of the
same period, fossil primates are notable by their absence, suggesting that at that time
South Africa lay outside the taxonomic diversity envelope for primates Other
mammal groups, such as ruminants and suids which occur in South African nliddle
Miocene deposits, tend to be lower in diversity compared with contemporary East
African and European forms. This could be because South Africa was near the
southern end of the diversity envelopes for these groups.
If the evidence is viewed at a parochial scale, only local or regional possibilities

can emerge from the data base. It is, therefore, necessary to keep in mind the scale
effect, by which one is enabled to see different patterns in the biosphere at different
scales. Confusing the scales may even yield conflicting results.
428
JNPLICATIONS OF PLATE TECTONICS
Many of the above possibilities could only occur if there were free access by
which organisms could expand and contract their geographic ranges between neighbor-
ing continents. During most of the Oligocene, Africa was so far removed from Europe
that few African lineages could colonize Europe. However, during this same period,
numerous lineages of Eurasian organisms managed to colonize Africa (Pickford,
1986a). This one-way traffic is perhaps best explained as the result of two factors:
a) the direction of ocean currents in the Tethys Seaway, and b) Wallace's Rule.
During the Oligocene, the Tethys Sea connected the Atlantic and Indian Oceans
permitting circulation of waters between the two oceans. It is postulated that circu-
lation was from east to west, and that there was a persistent component of current
toward the south (figure 16). This configuration would promote a southwesterly trans-
port of organisms from Eurasia to Africa at the same time that it hindered traffic in
the opposite direction.
It is also probable that Wallace's Rule was acting during the Paleogene just as it
does today, and that African mammals traversing the Tethys Seaway northward would
be subjected to the latitudinal effects by which taxonomic diversity decreases as
latitude increases. This double barrier, physical (Tethys water barrier) and ecological
(Wallace's Rule), may have acted in concert to prevent continental interchanges with a
northward component of movement. Conversely, organisms managing to cross the
Tethys Seaway southward from Eurasia to Africa would be entering parts of the
diversity envelope which would enhance their colonization and even promote increases
in diversity.
A comparable series of exchanges took place during the Plio-Pleistocene in the

New World. North American lineages colonizing South America often underwent
radiations once they arrived there (Marshall et al., 198Z, 1983). In contrast, lineages
which emanated from South America going northward seldom radiated to any great
extent or even became extinct after short tenures in the north.
DISPERSAL AND PLATE TECTONICS
During the Neogene in contrast, colonization of Eurasia by organisms of African

origin and vice versa was pervasive and frequent, and it appears likely that by the
beginning of the Neogene Africa was so close to Eurasia that the seaway separating
them was no longer an effective barrier to intercontinental faunal exchanges.
Eventually, by the beginning of the Orleanian land mammal age, Africa had made
definitive dry land contact with Eurasia, and many lineages of terrestrial organisms
were involved in interchanges between the two land masses (Pickford, 1981). This
quasi- or complete contact between Africa and Eurasia lessened or removed the
physical barrier to intercontinental faunal exchanges, but Wallace's Rule continued to
exert its influence on Neogene faunas, just as it still does with modern ones.
During much of the Miocene Epoch, however, the boundary between the
Palearctic and Ethiopian Realms appears to have been located considerably to the
north of its present position, and peri-Mediterranean areas of southern Europe as well
as much of southern Asia including most of Japan were essentially part of an expanded
tropical to subtropical belt, comprising enlarged Ethiopian and Oriental Realms
(Pickford, 1986b). It was this feature, more than any other, which led to the intro-
duction of numerous African mammals into southern Eurasia during the Miocene. Dry
land contact between the continents was an important, but not an essential, prerequi-
site for large scale interchanges to occur, but once such "dryshod" access had been
established, biogeographic constraints such as Wallace's Rule continued to regulate
faunal distribution patterns, just as they do today.
For example, despite the fact that the eastern half of the boundary between the
present-day· Pale arctic and Oriental Biogeographic Realms is thousands of kilometers
429
long and lacks any significant physical barriers, the faunas and floras have, at the
scale of the biogeographic realm, maintained their distinct distribution patterns.
However, during the Miocene, the introduction of organisms of African origin

into southern Eurasia involved numerous lineages of mammals, reptiles, and plants,
and it seems likely that, rather than all these groups adapting more or less simultane-
ously to Palearctic conditions, southern Eurasia merely became more tropical,
permitting many Ethiopian organisms to enlarge their distribution ranges northward
with a minimum of adaptation. In effect, southern Europe became sub-tropical and
therefore belonged to the northern part of the Ethiopian Biogeographic Realm. At the
end of the Miocene, conditions changed, with the result that the boundary of the
Ethiopian Realm shifted back southward to such an extent that North Africa became
part of the Palearctic Realm. This southerly shift led to the local extinction in
southern Europe of those organisms adapted to an Ethiopian Biogeographic regime, and
to its recolonization by Palearctic lineages. The same process took place in North
Africa, despite the existence of the Mediterranean barrier, with the result that the
Maghreb, which, during the Miocene, used to contain typical members of the Ethiopian
Realm, is nowadays dominated by Palearctic faunal and floral elements.
The main problem is to determine whether these north-south fluctuations in

distribution patterns were caused by inflation-deflation of the taxonomic diversity
envelopes, or by other regional or local causes, possibly including shifts in the position
of the equator. The solution to this problem cannot be found by recourse to parochial
evidence. The only possibility of obtaining answers is to examine a wide range of
faunas from various latitudes and longitudes on both sides of the equator.
THE DRIFT COMPONENT OF FLUCTUATING BIOGEOGRAPMC BOUNDARIES
If the concept of fluctuating biogeographic boundaries is correct, then it would

be interesting to determine what happened to cause such shifts. A possible explana-
tion is that the Miocene equator lay further north than it does today as suggested by
Koeniguer (1987) and Termier and Termier (1986). Northward drift of Africa toward
Europe could partly account for this apparent northward displacement of the
equator. It can be envisaged that evidence upon which one can reconstruct the
position of the equator became fossilized in sediments which were then passively
transported northward during plate tectonic processes, taking their information
content with them. However, the earliest discoveries suggesting a northward position
of the equator during the Miocene come from marine organisms, including sponges
(Termier and Termier, 1986) and mangroves (Koeniguer, 1987; Karyu et al., 1984),
which grew along the northern shores of the Mediterranean portion of the Tethys and
far to the east in Japan. In addition, many of the terrestrial organisms which suggest
the existence of tropical conditions in southern Europe during the Miocene, including
palms (Chepalyga, 1985) (figure 13) also provide evidence for the northward displace-
ment of the Miocene boundary between the Palearctic and Ethiopian Biogeographic
Realms. It is concluded, therefore, that far from continental drift, on its own,
accounting for a northward displacement on the tropical belt, and by inference, the
equator, there must be some other explanation to account for this pattern, if
Koeniguer's and the Termiers's observations are correct.
POLAR SHIFT AND CHANGES IN BIOGEOGRAPMC BOUNDARIES
If, as hypothesized by Koeniguer (1987) and Termier and Termier (1986), the
equator really was further north in the Old World during the Miocene, a suggestion
made independently on the basis of diverse lines of evidence, then one would expect
that it would be located further south in the New World than it is now. This, of
course, is because the equator is a great circle, and it would have to twist about an
axis rather than be displaced parallel to itself. Evidence from the New World could
support this contention, since there is a long-standing body of evidence indicating that
the southern tip of Patagonia (S0°S) was considerably more tropical during the
430
Miocene than it is now (Marshall et al., 198Z). Miocene faunas from North America, in
contrast, are apparently rather monotonous and of moderate diversity throughout the
Miocene.
Previous explanations for latitudinal shift in faunal and floral boundaries in

South America invoked the rise of the Andes as the agent causing an equatorward
shrinking of the South American tropics. However, they are unlikely to account for all
the observed changes, since they fail to take into account evidence from other parts
of the globe. This is not to say that the rise of the Andes had no effect on continental
climate of South America; undoubtedly it did. However, their growth is not sufficient
to cause such spectacular changes in the disposition of the tropical belt around the
globe. For such a thing to happen, a global or astronomical event presumably had to
occur.
In the Old World, evidence from the southern part of Africa indicates that it was
less tropical during the Miocene than it is now, which could be one reason why several
groups of tropical organisms, including primates, are lacking in its Miocene fossil
record. This evidence also supports the idea that the tropical belt was further north in
Africa during the Miocene than it is now.
If the equator occupied a different position or positions on the globe during the
Miocene, it follows that the poles of rotation of the globe would have been in different
positions from those that they occupy now, since pole position defines the equator, and
vice versa. Preliminary estimates based on the data that I have seen suggest that the
Miocene North Pole might have shifted about Z0° south of its present position, some-
where in the vicinity of the North Slopes of Alaska (figures 14-16) and approximately
the same distance from the North Magnetic Pole as the present rotational pole is.
Before accepting this concept we should ask ourselves whether there are any
tests that we can perform to refute the hypothesis of changed pole positions.
Fig. 15. Reconstruction of Miocene Biogeographic Realms

on the basis of distribution patterns of organisms
which originated in Africa and colonized large
parts of Eurasia. Note that in this model the
boundary between the Ethiopian Realm (small dots)
and the Oriental Realm (large dots) is gradational,
and a case might be made for considering them to
be parts of a single widespread biogeographic realm
with a comparable longitudinal extent to that of
the modern Palearctic Realm to the north.
431
Fig. 16. Maps contrasting the different histories of organismal
exchange between Africa and Eurasia during the Paleogene
and Neogene. Between the lower Oligocene and lower
Miocene, numerous Eurasian lineages colonized Africa {arrows
with tails) whereas only one mammalian lineage managed to
cross the Tethys in a northward direction. It is suggested that
the ocean currents flowing through the Tethys were from east
to west, and that there was a persistent southerly component
to the currents. This would facilitate transfers in a southerly
direction and hinder those in a northward direction. Modern
ocean currents in the Arafara Sea between Australia and the
Indonesian Archipelago provide a modern analogue, the main
difference being that in this case there is a persistent
northerly component to the currents. This has facilitated the
transfer of marsupials northwestward, but has hindered the
colonization of Australia by placentals.
During the Miocene, in contrast, Africa was so close to

Eurasia, and even in contact with it, that numerous terrestrial
lineages could make the exchange in both directions. How-
ever, Wallace's Rule continued to play its role throughout the
Miocene Period, with the result that the Palearctic and
Ethiopian Realms maintained their distinctiveness, only a few
lineages being able to make the necessary adaptive changes
permitting them to cross the major latitudinal ecological
boundary between the two realms. The colonization of south-
ern Europe by many lineages of African origin is perhaps best
explained by invoking the model in which the Miocene tropical
belt was further north than it is now.
432
TAXONOMIC DIVERSITY AND POLE LOCA'I10N
Work being done during the late 1950s and early 1960s dealt with taxonomic
diversity gradients and their potential for locating ancient pole positions (Stehli,
1968). This approach lost most of its momentum when the theory of plate tectonics
crystallized early in the '60s. The reason was that the plate tectonic model was felt
to explain the data in a more satisfactory way than did the polar wander model being
investigated at that time. This is undoubtedly true for the Paleozoic and Mesozoic
because the amplitude of continental drift that has occurred since those eras is so
great that any "signals" from taxonomic diversity patterns that may have existed have
been completely swamped by the "noise" yielded by subsequent drift. For the
Cenozoic, and specifically the Neogene, however, the amplitude of continental drift
has been small enough that taxonomic diversity signals are still clearly visible above
the noise introduced by drift.
Many groups of organisms can be utilized in studies of ancient latitudinal

taxonomic diversity, provided that their fossil record is complete enough. One such
group is the primates (Pickford, 1986c). Today most primates, apart from macaques
and one or two colobine species, are confined to the tropics (Pickford, 1987a). Within
the tropics, there is a marked diversity peak near the equator, which drops rapidly at
latitudes higher than 10° either side of the equator (Pickford, 1987a) (figure 17).
During the early Miocene, catarrhines were very diverse in East Africa (11 to 1Z
species known) but have never been found in European deposits older than MN4
(Pickford, 1986c). By middle Miocene times, however, the diversity of catarrhines in
East Africa began to decrease (six species at Maboko, dropping to three species at
Ngorora) (Pickford, 1981) whereas their diversity in southern Europe began to increase
(figure 18). One explanation that offers itself is that the northward shift in the
position of the tropical belt induced a comparable shift in the position of the latitu-
dinal taxonomic diversity peak (figure 6). In terms of present-day geography, this
Cercopithecidae Gorillidae
Fig. 17. Taxonomic diversity of extant African anthropoids. The highest diversity
levels occur near the equator and there is a rapid drop in diversity from
about 10° latitude either side of the equator. There is a lesser east-west
diversity change, principally a drop in East Africa with respect to West
and Central Africa, due to the semiarid to arid belt of country in the far
east of the continent.
433
EAST AFRICA EUROPE NWINDIA
Pliopiths
Numbers of species
Fig. 18. Changes in taxonomic diversity of Old World

anthropoids during the Tertiary and
Quaternary. A particularly noticeable
feature of the tropical African higher primate
fossil record is the marked diversity trough
which occurred during the latter part of the
middle Miocene and the late Miocene. In
contrast, the middle Miocene was a time
during which taxonomic diversity of higher
primates increased in southern Europe, a
diversity peak occurring during the middle
Miocene. Similarly, a diversity increase
occurred in northwest India during the middle
and late Miocene, followed by a decrease in
the latest Miocene. The shift of diversity
peaks from tropical Africa toward southern
Europe suggests that the tropical zone was
further north during the middle Miocene than
it was in the early Miocene and at the present
day.
434
would mean that East Africa would have been near the southern end of the diversity
gradient envelope, whil~ Europe would have been near its northern end. The peak
would have lain about ZO north of where it does now.
In late Miocene times, catarrhine diversity in Europe dropped, leaving only

macaques and a colobine in the continent (Ardito and Mottura, 1987). Meanwhile, in
the Plio-Pleistocene of East Africa, catarrhine diversity was increasing dramatically
with major increments taking place from about 4 Ma onward (Picford, 1987b). It is
possible that the posjtion of the tropical belt was shifting southward again, taking the
taxonomic diversity envelope with it, and that since about 3 Ma, it has occupied more
or less the same position that it does today.
Examination of the South American fossil and present-day evidence yields data
of a complementary nature. For example, primates which inhabited Patagonia, about
S0°S latitude during the middle Miocene, were relatively abundant elements of the
faunas. Nowadays, no primates occur in Patagonia, nor for that matter within much
of Argentina to the north, their most southerly present-day distribution being about
30°S (figure lZ). The present-day South American· primate diversity peak is near the
equator, whereas during the Miocene it seems to have been so far south that the limbs
of the diversity envelope almost reached the sguthern tip of the continent (figure 7).
A shift in position of the tropical belt by 15-ZO to the south would satisfy the data, a
figure which accords with preliminary estimates of its postulated northward shift in
the Old World.
If this scenario is valid, then the apparent northward and southward fluctuations
of the boundaries of biogeographic realms in the Old World become more simple to
understand. A northward shift in Old World biogeographic boundaries would lead to
northward shifts in the distribution ranges of Ethiopian organisms into areas where
they previously could not survive. A retum southward of the biogeographic boundaries
would lead to a retreat of the tropical to sub-tropical faunal elements typical of the
Ethiopian Realm from areas which had reverted to their former Palearctic status. If
this is what happened, then I consider that we possess a dynamic model that can
explain many of the features of the European fossil record which form the basis for
European Neogene biostratigraphy.
AFRICAN CONTRIBUTIONS TO EUROPEAN BIOSTRATIGRAPHY
Into the latter category fall the successive arrivals into southern Europe of taxa
considered to have had an African origin, including Brachyodus, gomphotheres,
deinotheres, creodonts, hominoids, hyracoids, giraffids, tubulidentates, cercopithe-
coids, hippopotamids, and elephantids, among other mammalian lineages. Since these
taxa are readily identified and easily distinguished from authochthonous European
lineages, they are frequently given prominence in definitions of European land
mammal chronology.
The Hipparion datum represents a comparable colonization of the Old World by a

taxon with no known Old World ancestry, which was an immigrant from the New
World. It is, therefore, accorded a great degree of confidence in Old World biostra-
tigraphic schemes, its earliest appearance in Eurasia and Africa generally being
employed to define the base of the late Miocene Period in continental sequences of
the Old World. An important source of this confidence is the ease with which it can
be correctly identified at the generic level, and its abundance in the fossil record.
The finding of even a small fragment of Hipparion tooth in an Old World sedimentary
unit is sufficient to reveal that the strata must be late Miocene or younger in age.
The equids Anchitherium an'\sfquus provide equally valuable biostratigraphic data for
the middle Miocene and Plio- etstocene, respectively.
From the point of view of Africa's contribution to the European fossil record, its
influence cannot be overestimated. Indeed, several important biostratigraphic
markers entered southern Europe from Africa, some of them being so useful that
435
European land mammal ages are defined on the basis of their appearance in the
southern European fossil record. Among these are the appearance of Brachyodus at
the continental equivalent of the Oligo-Miocene boundary, and the influx of
Proboscidea at the beginning MN4a. I think that it is likely that further research
along these avenues will lead to more precision in European large mammal biostratig-
raphy. It is already evident that the influx of Hippopotamus in the latest Miocene is a
useful biostratigraphic event, and there were other groups, such as hyracoids,
tubulidentates, hominoids, and cercopithecoids (colobines and macaques), which are of
equal potential.
What is required is a more concerted effort to compile a complete list of all

sites in Europe which have yielded taxa of African origin. There is much information
already in the literature, but there is probably a great deal more unpublished data
sitting unnoticed in museum drawers. Many of the fossils might be insignificant from
the point of view of increasing knowledge about the anatomy of the groups, but each
one, no matter how incomplete, can yield information of potential use in biogeog-
raphy.
EURASIA'S CONTRIBUTION TO AFRICAN BIOSTRATIGRAPHY
Equally important from an African biostratigraphic perspective are the introduc-

tions of Eurasian faunal elements into Africa. For example, the suid Listriodon makes
its appearance abruptly in the East African fossil record about 16 Ma, presumably
from Eurasia, since none of the known prior African suid lineages are suitable
ancestors for it. Likewise, the arrival of tetraconodonts in the middle Miocene,
Hipparion in the late Miocene, Suini in the early Pliocene, Equus in the late Pliocene,
and Camelus in the early Pleistocene are reliable key biostratigraphic events in
tropical Africa, and there are several others, including Sivameryx (early Miocene),
Ancylotherium (late Miocene), Hystrix (latest Miocene), and Lepus (latest Miocene).
Important African biostratigraphic markers based on taxa incoming from Eurasia are
not confined to mammals. For example, the bivalve Corbicula makes its appearance
in African deposits about 3.4 Ma and is common in lacustrine strata younger than this.
It is evident to me that a more complete understanding of the development of

the Ethiopian faunas during the Neogene will automatically lead to a better under-
standing of the European sequence. With this in mind, I have been compiling as
complete a list as possible of large mammals from as many sites in Africa that I know
about, with the aim of determining the evolution of the African faunas. An obvious
starting point is the East African sequence, which is reasonably well calibrated by
radioisotopic age determinations. In the future it will be necessary to examine the
faunas from South Africa and from the northern margins of the continent in order to
fill out the faunal history on a continental basis. This will facilitate correlations with
Europe, as well as permitting reconstructions of diversity gradient envelopes and
identification of major faunal events. At present African micromammals are. less
useful for intercontinental biostratigraphy than large and medium sized mammals for
three reasons. Firstly, their African fossil record is replete with gaps which makes it
difficult to understand the timing of the changes which occurred. Secondly, rodents
generally have restricted geographic ranges, with the result that tropical Africa and
Europe have had few, if any, rodent species or genera in common, even during the
Miocene. Thirdly, many rodent taxa apparently evolved more slowly in the tropics
than did the large mammals, although why this should have been so is an unresolved
paleontological issue.
TAXONOMIC DIVERSITY CHANGES IN THE

PLIQ-PLEISTOCENE. OF TROPICAL AFRICA
Study of the East African Plio-Pleistocene mammal record from the point of
view of diversity (figure 19) reveals that the diversity of fossil species fluctuated
widely. Such fluctuations are sometimes considered to be an artifact caused by the
436
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Fig. 19. Fluctuations in taxonomic diversity of selected mammalian groups in the east African
Plio-Pleistocene sequence. Dotted portion of graphs represent levels which exceed
the modern diversity of the lineages, which is represented by the unshaded portion of
each graph. There is a common theme through time in the sequence whereby peaks
and troughs in diversity of several groups coincide in time (peaks at 5.5, 4.0, Z.9, and
.,.w Z.3 Ma and marked troughs at 3.6, Z.6, and 1.8 Ma) •
.....
numerous taphonomic biases which acted between the paleobiosphere and the collector
of fossils, and, as such, examination of ancient diversity has not been a popular subject
for research. However, parts of the East African fossil record are so complete that
ancient diversity levels at certain horizons are greater than diversity in the same
regions today. If there is a pattern of under-representation in the fossil record, then
it is important to understand why during some periods in the past diversity was even
greater than it is today. Furthermore, it is clear from figure 18 that increases and
decreases in taxonomic diversity in several groups of mammals occurred simultane-
ously. For example, major diversity peaks occurred about Z.9 and Z.3 m.y. ago in East
Africa; peaks which were preceded by, separated from, and followed by periods during
which diversity was less than it is today. Less readily perceivable diversity peaks
occurred about 5.5 and about 4.0 Ma.
My first attempt to explain these fluctuations is represented in figure ZO. I

thought it likely that biota in the Omo Basin, from which the basic data comes,
experienced changes in climate and vegetation cover (for which there is independent
evidence) from an initially humid ecosystem (A in figure ZO) in which a fauna of
moderate diversity occurred (3.5 to 3.0 Ma). With the change in climate, a sub-humid
ecosystem became established in the area with its own mammal fauna (B in figure ZO),
but during the switch over, when patches of both ecosystems were present in the
basin, the mammal fauna appeared to undergo a short-lived increase in diversity.
2m.y.
Fig. ZO. Ecological interpretation of the data of figure

16 considered as a local phenomenon. In this
probably incorrect model, changing environ-
mental conditions are visualized as having led
to short-term increases in diversity as the
result of the establishment of mosaic habitats
in East Africa, each part of the mosaic con-
tributing to a diversity total which is greater
than the usual diversity found in more
uniform ecosystems. It appears, however,
that this model breaks down when examined
from a continental perspective, because
comparable changes seem to have occurred at
the same time in Southern Africa and in
Europe.
Note that, in the Omo Basin, hominid diver-

sity changed twice at precisely the same time
that changes occurred in other mammalian
groups.
438
About z.s Ma the climate changed again, leading to the establishment of a semiarid
ecosystem (C in figure ZO), again with a short period when a mosaic of environmental
conditions existed in the area, represented by a short period during which faunal
diversity apparently increased. It was noted that hominid diversity also increased and
decreased at these times (lower part of figure ZO).
However, far from being a localized phenomenon restricted to the Omo Basin,
such diversity fluctuations appear to have occurred on a much wider scale, with
comparable fluctuations in southern Africa and Europe. It now seems more likely to
me that we have evidence for changes in diversity caused by some global scale factor,
such as fluctuations in solar energy capture. If this is so, then one could expect
similar diversity changes to have occurred in many places in the globe, and examina-
tion of suitable fossil sequences is recommended.
In this model, diversity increase refers principally to a phylogenetic radiation

rather than to a shift in distribution patterns of existing faunas, whereas a decrease in
diversity refers predominantly to an extinction event, rather than to emigration of
taxa from a region. In order to test this hypothesis, I examined the morphological
variability of taxa during the same period of time in the same basin, since phyletic
evolution should be recorded in the fossils as changes in morphology or size through
time.
FLUCTUATIONS IN VARIABILll'Y IN EAST AFRICAN

PUQ-PLEISTOCENE MAMMALS
Examination of variability in several groups of Plio-Pleistocene mammals of

tropical Africa (figure Zl) shows that there were periods during which variability
within lineages was relatively constant, interspersed by short periods during which the
coefficients of variation increased markedly. For example, in the proboscidean
lineage (Elephas recki), variability in molar plate height and enamel thickness was
relatively constant, even though the mean was drifting slightly over time, but for
short periods (e.g., about Z.4 Ma) the variablity was about twice as great as usual. In
the suid lineages Mesochoerus and Notochoerus, the coefficient of variation in third
molar length increased dramatically about z. 7-Z.S and Z.4-Z.3 Ma, whereas "normal"
coefficients of variation occurred prior to these periods, between them, and after
them. These short periods during which variability was greater than usual coincide
closely in time with periods during which taxonomic diversity was higher than usual.
Hominid diversity in the Omo Basin underwent increments about z.s and Z.4 Ma,
precisely at the time that other mammalian lineages were diversifying. It is perhaps
no mere coincidence that these events occurred together and it seems likely that they
all represent tandem responses by different mammalian groups to changes in environ-
ment.
Such contemporary responses to environmental change by numerous groups of

organisms comprise the basis for "tandem evolution." It is interesting to speculate
that hominids of the Plio-Pleiatocene were probably no different from other mammals
in this respect, and that they also experienced radiations and extinctions at the same
times that other groups were affected.
CONCLUSIONS
I would like to emphasize that an understanding of the mechanisms which cause

changes in the distribution patterns and variability of organisms will increasingly be
seen as an essential field of study, without which disciplines like biostratigraphy will
remain merely descriptive pursuits. Whether the models of fluctuating biogeographic
boundaries, polar shift, latitudinal taxonomic diversity gradients, and fluctuating solar
energy capture models are correct or not does not affect the utility and practice of
biostratigaphy, but they certainly lead us to ponder about how the observed faunal
changes may have been caused.
439
Plote height Enomel thickness
10 , .... , ; l i ~ : "P'o
my
K 1·5
J7
H4
H2
H
.....
''
i
,-
Period of continued
2
;:: ''
'
I
G
F
F4
E
0
(4 ::3
(
of ·
3·5
A .,"'2
0
-5
20
z
M 4
u
Fig. Zl. Changes in morphological variability in various mammalian lineages. In the

Omo succession, the coefficient of variation in various measurable param-
eters of the molars of proboscideans and suids varied through time. Apart
from a shift in the value of the mean over time, it is apparent that there
were at least two short periods during which the coefficient of variation
was greater than "usual" in several mamma.l groups. These periods of
"excess" variability coincide in time with periods of increased taxonomic
diversity (see figure 17). The diversity of hominid species in the basin
changed during these same periods.
Such tandem evolution demands an evolutionary model in which selection is

not only contemporary, but acts in parallel or convergent ways in phylo-
genetically distinct lineages. This in turn suggests that exogenetic factors
are at work. Since variability and taxonomic diversity fluctuations occurred
contemporaneously, one possible explanation is that the quantity of solar
energy captured by the globe fluctuated over time and that peaks in divers-
ity and variabilty coincide with increased solar energy input into the globe,
whereas diversity troughs and decreases in variability correlate with
decreases in solar energy capture. (Basic data from Beden, 1979; Cooke,
1976.)
440
ACKNOWLEDGMENTS
I would like to thank the organizers of the NATO Advanced Research Workshop
for the invitation to participate in such a stimulating workshop. I am also anxious to
thank Professor A. B. Chiarelli for support and encouragement.
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Beden, M., 1979. Les elephants (Loxodonta et Elephas) d'Afrique Orientale:
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(1), p. 1-ZZ4; (Z), p. ZZ5-567.
Broin, F. de, 1987. Les tortues et le Gondwana. Bull. Groupe Fran~ais d'etude du
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Chepalyga, A.L., 1985. Climatic and eustatic fluctuations in the Paratethys Basins
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Cooke, H.B.S., 1976. Suidae from Plio-Pleistocene strata of the Rudolf Basin, in
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Karyu, T., Itoigawa, J., and Yamanoi, T., 1984. On the middle Miocene palaeo-
environment of Japan with special reference to the ancient mangrove swamps, in
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Lavallard, J-L., 1988. Pourquoi la Terre tourne moins vite. Sciences et Avenir, v.
494, p. 34-39.
Malingreau, J-P. and Tucker, C.J., 1987. La vegetation vue de l'espace. La
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Marshall, L., Webb, S., Sepkoski, J ., and Raup, D., 198Z. Mammalian evolution and the
Great American Interchange. Science, v. Z15, p. 1351-1357.
Marshall, L., Hoffstetter, R., and Pascaul, R., 1983. Mammals and stratigraphy:
Geochronology of the continental mammal-bearing Tertiary of South America.
Palaeovertebrata Mem. extraord. 1983, p. 1::;~·
Miller, K. and Fairbanks, R., 1985. Cainozoic o 0 record of climate and sea level.
S. Afr. J. Sci., v. 81, p. Z48-Z49.
Pickford, M., 1981. Preliminary Miocene mammalian biostratigraphy for western
Kenya. Jl. Human Evol., v. 10, p. 73-97.
Pickford, M., 1986a. Premiere decouverte d'un hyracoide paleogene en Eurasie. C. R.
Acad. Sc., Paris, v. 303, p. 1Z51-1Z54.
Pickford, M., 1986b. Geochronology of the Hominoidea: A summary, in Else, J.G. and
Lee, P.C. (eds.), "Primate Evolution." Cambridge Univ. Press-;-Cambridge, p.
1Z3-1Z8.
Pickford, M., 1986c. The geochronology of Miocene higher primate faunas of East
Africa, in Else, J.G. and Lee, P.C. (eds.), "Primate Evolution." Cambridge Univ.
Press, Cambridge, p. 19-45.
Pickford, M., 1987a. The diversity, zoogeography and geochronology of monkeys.
Human Evol., v. Z, p. 71-89.
Pickford, M., 1987b. The chronology of the Cercopithecoidea of East Africa. Human
Evol., v. Z, P• 1-17.
Pickford, M., 1987c. Concordance entre la paleontologie continentale de l'Est
Africain et le!! evenements paleo-oceanographiques au N~og~ne. C. R. Acad. Sc.
Paris, v. 304, p. 675-678.
Stehli, G., 1968. Taxonomic diversity gradients in pole location: The recent model, in
Drake, E.T. (ed.), "Evolution and Environment." Yale Univ. Press, New Haven
and London, p. 163-Z19•
441
Termier, H., and Termier, G., 1986. Relations des spongiaires avec la Tethys. Bull.
Groupe Fran~ais d.'etude du Gondwana, v. 9, p. 45-67.
Tucker, C.J., Fung, I.Y., Keeling, C.D., and Gammon, R.H., 1986. Relationship
between atmospheric COz variations and a satellite-derived vegetation index.
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Wallace, A.R., 1876. The Geographical Distribution of Animals. Macmillan, London.
442
MIOCENE PALEOECOLOGY OF PA~ALAR, TURKEY
Bema Alpagut
University of Ankara
Turkey
Peter Andrews
British Museum (Natural History)

London, England
Lawrence Martin
State University of New York

Stony Brook, New York, U.S.A.
INTRODUCTION
The study of mammalian faunas can be said to have three aims. One is to use
the evidence of change through time to investigate evolutionary processes. The
understanding of such processes is clearly fundamental to the interpretation of both
living and past faunas. A second aim is to use similar evidence of change through time
to investigate the emergence and diversification of particular lineages, and this can be
used both for the better understanding of the groups in question and for the correla-
tion of faunas between sites. Relative time periods may be defined on the basis of
certain marker fossils, and these may be taken as representing evolutionary stages
throughout a region. The third aim is to use the types of animal represented in a
fauna to indicate paleoecology. The community structure of the faunas can be com-
pared with those of extant faunas, providing the basis for the interpretation of ecolog-
ical change through time and between regions.
The first of these aims does not concern us here. The second aim also can be
mentioned only very briefly. Our main purpose in carrying out fieldwork in Turkey is
to increase our sample of hominoid primates with the purpose of coming to a better
understanding of the evolution of the sivapithecine hominoids. Our site at Pa~alar
(figure 1) has yielded more than 500 specimens of hominoid a.ssigned at present to two
species (Tekkaya, 1974; Andrews and Tobien, 1977). These have morphological simi-
larities to later Miocene species of Sivapithecus (such as thick molar enamel, incisor
heteromorphy, and flattened enamel-dentine junctions), which probably indicate
phylogenetic affinity, but they differ in the retention of primitive characters of the
teeth, such as molar cingula and incisor medial pillars. These are character states
present in early Miocene African hominoids but lost in later Miocene sivapithecines.
The combination of advanced sivapithecine characters with primitive hominoid reten-
tions is seen in the hominoids from just three sites: P~alar and <;andir in Turkey, and
Neudorf an der March in Czechoslovakia. In other aspects of their faunas these sites
show great similarity as well, and it is likely that they are similar in age and paleo-
ecology. The comparative approach clearly has much to offer in the interpretation of
these faunas.

t
IZMIR
................5 10 km
~_....
i
Fig. 1. Locality maps: above, westem Anatolia showing loca-
tion of Pa,alar; below, the position of the site (bottom
middle) and Neogene sediments (n) in an embayment of
Mesozoic rocks (hatched), while further to the north the
Neogene deposits are covered by Quatemary sediments
(Q).
444
Our work on the Pa~alar hominoids will be described in a separate publication,
and in the present paper it is our intention to concentrate on aspects of the paleoecol-
ogy of the Pa;salar fauna. This is the third of the aims of faunal analysis described
above, and in concentrating on the paleoecology of a single site we will be able to go
into some detail on the relevance of paleoecological work and the significance of
taphonomy for paleoecologia! inferences. In describing any mammalian fauna, it is
necessary to investigate the manner in which it has been accumulated before drawing
any paleoecological conclusions from it, for it is commonly the case that fossil faunas
are selectively biased by whatever agents are acting to accumulate and preserve
them. Predator assemblages, for instance, reflect the tastes and hunting skills of the
predator, which selects from the animals available to it only those that it can kill and
eat. Water-transported assemblages equally reflect the movability of different bones
in terms of their size and shape, and the end result may be very different from
primary bone source(s). An understanding of these taphonomic events may go some
way towards making some allowance for observed biases, and this can only be achieved
by comparison with taphonomic agents acting today. Similarly, the study of paleo-
ecology relies ultimately on knowledge of present-day ecology. The approach that we
shall use will be to analyze the community structure of entire faunas (Andrews et al.,
1979).
Pa;salar is a site singularly well suited to paleoecological analysis, because the

accumulation of the fossiliferous sediments can be shown to have been very rapid and
to have been derived from local sources. In the next section the stratigraphy of the
site will be described briefly, and this will be followed by an account of the fauna,
encompassing both the species represented and any information on their taphonomy.
Based on the results of the faunal descriptions, the paleoecology of the site will then
be described.
THE SITE AT PA\iALAR
The site at Pavalar was discovered during the Turkish-German lignite survey of
Turkey between 1965 and 1969 (Sickenberg, 1975). Early excavations were directed by
Heinz Tobien in 1969 and 1970 (Sickenberg, 1975; Andrews and Tobien, 1977). After
1983, excavations were restarted by us on behalf of the Ministry of Culture and
Tourism, Department of Antiquities, and the University of Ankara, Department of
Paleoanthropology (Alpagut, 1984, 1985, 1986, 1987, 1988), and excavations are still
continuing.
The fossiliferous sediments 1 km to the south of Patalar village outcrop on the

edge of the Gonen basin (figure 1). The fauna from a single sedimentary unit will be
described here, an unsorted gravelly sand near the base of the Pa~alar sequence (figure
Z). Several additional units of fine sands and sandy silts are associated with the top
and bottom of the unsorted sands, but these are unfossiliferous for the most part and
will not be considered here. In any event, the bones from the unsorted gravelly sands
make up over 1).5% of the identifiable faunal remains from Patalar.
The unsorted sands were deposited as an unsorted debris flow off the adjacent
hillslopes. The main massif of the hills to the south and west of the site consists of
intrusive granites, but outcropping immediately to the west, and underlying the
Miocene sediments, is a folded series of Mesozoic rocks consisting of the P~alar
marbles and the K1z!ltepe metamorphic series. These form the lower slopes of the
hills next to the site, and they also make up the source rock from which the sediments
were derived. Hardly any granite has been found in the sediments, and the major
source rocks are the K1z1ltepe metamorphics. From this it is evident that the source
area of the Patalar sediments was from a maximum of Z-3 km from the site.
The sediments making up the gravelly sands are largely unsorted, but there is
evidence of fining of clasts up the sequence. Clasts up to the size of pebbles are
found throughout, but whereas 70% of the lower deposits consist of coarse sand,
gravel, and pebbles; this is reduced to 40% at the top, with another 40% medium and
445
fine sand and the remaining ZO% silt and clay. In the main part of the exposure the
change is gradual, and there is no evidence of any break in sedimentation. The lack of
any evidence of bedding or other structures indicative of water action together with
the lack of sorting suggests rapid deposition as a debris flow for this part of the
deposit. Certain of the features such as vertical gravel pipes, extend through the
vertical extent of the sands (figure Z). The limits of these pipes are discrete, and
since there is no sign of horizontal stratification within them it is concluded that they
represent single depositional episodes. There is no evidence that they are secondary
features; on the contrary, they are associated with potholes cut into the underlying
silt, and bone orientations in the surrounding sands are related to the gravel pipes. It
is concluded from this that the main part of the unsorted gravelly sands was deposited
as a single depositional event. This is envisaged as the result of a massive flood, first
producing the scouring evident in the underlying deposits and followed by massive
outwash of unsorted debris from the adjacent hillslopes.
The bones present in the unsorted sands are scattered throughout the deposit
approximately in proportion to the variations in particle size. Small mammals are
mainly concentrated in the areas of gravelly fine sand in the upper parts of the unit,
being rare in areas of coarse sand and almost absent in the pebble and gravel lenses.
Larger mammals are present but sporadic in the finer grades of sand, more abundant
in the coarse sands, and locally very abundant in the pebble and gravel lenses. This
distribution suggests that the bones form part of the sediment load and were carried
to the site together with the sediments. By the same token, they were therefore
derived originally from the same location as the sediments, so that it may be inferred
that they were local in origin, and accumulated very rapidly, as were the sediments.
This tells us two things about the animals in the Pa~alar deposits; they lived close to
the present site, probably mostly on the hillslopes within Z-3 km of the site, and they
were ecologically contemporaneous. The second conclusion stems from the rapidity of
deposition indicated by the geological evidence, which suggests the unsorted sands
could be the resuJ.t of a single storm.
Fig. z. General view of the site at Pa~jalar near the end of the
1987 season. The three main sedimentary units are
numbered 1, the unfossiliferous lower calcareous silt, Z,
the unsorted gravelly sand which is the main fossiliferous
unit, and 3, the upper calcareous silt, which is poorly
fossilif~rous. The top of the lower calcareous silt is
irregular, with many potholes, one of which is arrowed in
this figure.
446
THE PA~ALAR FAUNA
Descriptions of the faunal remains from Pa1ialar are not yet complete. The
following summary is therefore an interim account based in part on descriptions of the
earlier German collections (Sickenberg, 1975; Gasiry, 1976; Schmidt-Kittler, 1976;
Reissig, 1976; Andrews and Tobien, 1977; Hi.inermann, 1975, 1985; Engesser, 1980) and
in part on our unpublished collections (personal communications from M. Fortelius, A.
Gentry, and L. Jacobs).
Primates
Two species of primate have been described from Pa~alar by Andrews and Tobien
(1977). They were assigned to Sivapithecus darwini and R.amapithecus wickeri, large
and small species of thick enameled hominoid, respectively. The identification of the
large species is still accepted here, together with the similarity of the Pa~alar form
with that from Neudorf Sandberg in Czechoslovakia. It is a species the size of S.
sivalensis from the Siwaliks (Pilbeam et al., 1980), but in the retention of mol~
cingula and incisor morphology it preserves primitive hominoid characters lost by the
Siwalik species (and by other later Miocene taxa, for example, Dryopithecus).
The small species from Pa~alar was linked with the yandir mandible by Andrews
and Tobien (1977) in the genus Ramapithecus. Since that time, Ramapithecus has been
synonymized with Sivapithecus and the African species wickeri returned to its origi-
nally named genus, Kenyapithecus. While recognizing the great similarity between the
Pa1ialar species and the African one in their dentition, there are many details of the
mandible (from yandir), the lower canines and postcranial morphology (from Pa~alar),
and the lower third premolar (from both) that support generic distinction from
Kenyapithecus. We are therefore using the original attribution made by Tekkaya
(1974) for the <;andir mandible, Sivapithecus alpani, for both the yandir and smaller
Pa~alar specimens.
Two species of Sivapithecus are therefore recognized here, §_. darwini and §_.
alpani. There is some doubt, however, if these two species are actually distinct.
There is not space to go into this matter in any detail here, but there is little morpho-
logical difference between the two groups, other than incisor morphology, and the size
variation is comparable to that of the living orangutan. On the other hand, there is
little overlap in size between the two groups, and their separation is usually unambig-
uous, so that for the moment we are retaining the two species.
Sample sizes at the end of the 1987 season were 180 specimens of~· alpani and
Z79 specimens of§_. darwini. In addition, there are 10 unattributed postcranial frag-
ments. Only two maxillae and one mandible have been recovered during our excava-
tions, but there is abundant evidence that the isolated teeth that make up the bulk of
the collection belong to a limited number of individuals. It is possible to match wear
facets of upper with lower teeth and contiguous teeth along tooth rows, and one case
in particular has a number of teeth from one vertical gravel pipe fitting together.
This work is still in progress. Most of the teeth have their roots missing, and there is
some evidence of slight rounding of the bases of the crowns on about ZO% of the
.teeth. A very few show evidence of extensive rounding, while some, conversely, have
retained even fragile roots without any damage.
Jnsectivora
Five insectivore species have been described by Engesser (1980). By far the
most common species is Schizogalerix pasalarensis, the living counterparts of which
inhabit moist woodlands and in fact may be partly aquatic. There is an indeterminate
erinaceid and at least two soricids, but the remains of these animals consist only of
isolated teeth and they have yet to be identified. A few specimens of Desmanodon cf.
minor have been described by Engesser (1980).
447
It is apparent from the above that insectivores are poorly represented at
P~alar. We have found some postcrania of Desmanodon and one mandible of
Schizogalerix, but apart from this the remains consist entirely of isolated teeth.
These may be broken, with cusps knocked off and the roots damaged, but there is no
evidence of rolling of the specimens. A more detailed study is in preparation by
Jacobs.
Lagomorpha
Two taxa are present at Pasalar, Prolagus cf. oeningensis which is relatively
abundant, and a new species of Alloptox, which is less common. The former is repre-
sented by mandibles and teeth, the latter by isolated teeth only. In both cases there is
extensive damage to the surfaces of the crowns, with impact damage and other
evidence of attrition, but there is little evidence of rolling of the specimens. There is
no evidence of carnivore damage, either in the form of tooth marks or of digestion of
the enamel surfaces. A more detailed study is in preparation by ~evket ~en.
Rodentia
Eight rodent species have been recognized. The common species are the
cricetodontids Megacricetodon and Democricetodon, which have yet to be formally
identified. Turkomys pasalarensis has been described by Tobien (1978). These apart,
there are species of arboreal glirid, fossorial spalacid, identified as Pliospalax, and a
number of relatively rare taxa. A more detailed study is in preparation by Jacobs,
Flynn, and Engin Unay.
A small number of rodent incisors show signs of alteration on their enamel

surfaces as a result of digestion by a predator. Digestion removes and pits the enamel
to varying degrees depending on the nature of the predator and also on a number of
additional variables such as the age of the predator individual, time of year, and state
of hunger (Andrews, 1989). Five categories of predator modification have been recog-
nized for recent predators (Andrews, 1989). These are as follows:
Category 1 minimal digestion, molars Q-3%, incisors 0-13%

Category Z moderate digestion, molars 4-6%, incisors Z0-30%
Category 3 heavy digestion, molars 18-ZZ%, incisors 50-70%
Category 4 heavy digestion, molars 50-70%, incisors 60-80%
Category 5 extreme digestion, molars 50-100%, incisors 100%
Just over 1% of the PB.falar rodent incisors show evidence of digestion, which is at the
bottom of this range, but there is every likelihood that many of the rodents in the
Pqalar sample never passed through a predator at all but represent random death in
the cachement area of the deposits. The low percentage may therefore be misleading.
Degrees of alteration produced by digestion are closely correlated with numbers

affected in recent predator assemblages. Category 1 predators like Barn owls (Tyto
alba) and a number of other owl species produce only slight penetration of the enamel
during digestion, and where there is deeper penetration it is restricted to very local-
ized areas. Category Z predators like the Long-eared owl (Asio otus) and several other
owl species are similar except that deeper penetration of the enamel is more frequent
and is always restricted to the tips of the incisors. Category 3 predators like the
European eagle owl (Bubo bubo) and Tawny owl (Strix aluco) produce extensive altera-
tion of the enamel surface of rodent incisors, often with nearly complete removal of
enamel and some digestion of the dentine, producing a wavy outline to the tooth.
Category 4 predators like the Little owl (Athene noctua) and some diurnal raptors
produce very extensive enamel and dentine destruction, and Category 5 predators like
the red fox (Vulpes vulpes) and coyote (Canis latrans) are even more extreme, with
collapse and distortion of the teeth (Andrews, 1989). At Patalar, the only alteration
of the incisors seen is equivalent to Category 1, and no damage has been seen on any
of the molars, and this would suggest alteration by a Category 1 predator. This is
consistent with the low percentage of teeth affected by digestion present.
448
Even with this much information, it is not possible to identify the predator
producing the digestion of the P~alar teeth. Category 1 predators include mainly
specialized hunters (Andrews, 1989), so that predator bias is to be expected, particu-
larly against smaller or less common animals, but the exact'nature of this cannot yet
be identified. Other possible sources of small mammal bone from incidental deaths,
pitfalls, and accidental burials may further distort the proportions of species present
(Andrews, 1989).
Carnivora
Sixteen species of carnivore have been described from the Pallalar deposits by
Schmidt-Kittler (1976). This is almost certainly too many taxa, for it presupposes the
occurrence together of congeneric species of similar size and morphology. This is the
same problem that occurs for the two hominoid species. In living faunas in tropical
Africa, closely related primates tend to be allopatric, and where two congeneric
species do occupy the same area they have considerable behavioral and morphological
differences. This is even more true of carnivore species; small carnivores, for
instance, are either allopatric or have very distinctive adaptations, and the presence
of three species of a single genus, for example the three species attributed by
Schmidt-Kittler (1976) to Protictitherium, is not seen in living faunas. This problem
was recognized implicitly by Schmidt-Kittler, who stated that the nine arctoid species
(= Canoidea of Simpson, 1945) present at PC1falar, out of 16 carnivores, represented
too great a species diversity for the type of environment envisaged (see below). We
agree with this conclusion and would like to raise the possibility that the number of
carnivore species in the Pas>alar fauna has been unwittingly exaggerated. Having said
that, it is still the case that carnivores are very well represented at Pallalar. There
are primary felid predators, scavengers (assuming Percrocuta to be so), small mammal
predators, and the high diversity of omnivorous-type canoids. These correspond to the
full range of carnivore adaptations seen today in extant faunas.
Table 1. Pa~alar Faunal List
Sivapithecus darwini Protictitherium aff. gaillardi

Sivapithecus alpani Protictitherium new species
Soricid species A & B Percrocuta miocenica
Desmanodon cf. minor Percrocuta new species
Schizogalerix pasaJarensis Pseudaelurus cr. quadridentatus
Erinaceinae Pseudaelurus larteti
Prolagus cf. oeningensis Orycteropus
Alloptox cf. gobiensis Deinotherium giganteum
Turkomys pasalarensis Gomphotherium angustidens
Megacricetodon Pliohyrax
Democricetodon Anchitherium
Pseudodryomys Beliajevina tekkayai
Sciuridae Brachypotherium
Ctenodactylidae · Aceratherium
Pliospalax Chalicotherium
Rhizomys Listriodon cf. splendens
Amphicyon cf. major Listriodon sp. nov.
Hemicyon sansaniensis Conohyus simorrense
Plithocyon Taucanamo
Ursavus cf. primaevus Dorcatherium
Ursavus aff. intermedius Palaeomeryx
Plesiogulo Micromeryx
Proputorius Euprox
Trochictis Giraffokeryx
Lutrinae Caprotragoides
Protictitberium intermedium Kubanotragus
449
The most abundant of the carnivores in our Pa!jalar collections are the hyaenids
assigned to the genus Percrocuta. Viverrids show the greatest diversity and are also
common, but the other species of carnivore are all rare. All are mainly represented
by broken teeth, and in our excavations we have so far found only two more complete
specimens, single mandibles of Amphicyon and Protictitherium. Apart from the
breakage, the material is too fragmentary to provide much taphonomic information.
A more detailed study is in preparation by Mustafa Giirbuz.
Proboscidea
Two species of proboscidean are present in the P~alar deposits (Gaziry, 1976).
Deinotherium is uncommon; we have found a partial dentition and some isolated
molars in the course of our excavations. The common species is Gomphotherium
angustidens, again represented mainly by molars, but we have also found parts of two
skulls and a crushed but nearly complete humerus. Parts of one of the skulls are
dispersed vertically in one of the gravel pipes, with a lateral extent of less than half a
meter square and a vertical distribution of more than one meter. Many of the molars
are complete, but may be broken diagenetically. None of the tusks are complete,
although segments of up to 70 em are known, but some have been completely exploded
by diagenetic processes. The complete teeth do not show much evidence of transport,
but there are in addition many isolated fragments of proboscidean enamel that show
signs of transport in their rounded edges and chipped surfaces. These are found
completely isolated from other proboscidean remains, and undoubtedly they were
broken pre-depositionally. A more complete study is in preparation by Tobien.
Perissodactyla
Three groups of perissodactyl are present in the P~alar fauna. Equids are
represented by Anchitherium, a horse with low crowned teeth suitable for browsing
soft vegetation but not for grazing. It is less abundant than are several species of
bovid, giraffid, and suid, which will be described in the next section, and it is less
common than either of the two hominoids, but it is still one of the main herbivores at
the site. Chalicotheres are only represented by a few teeth. In contrast to this,
rhinocerotids are almost as common as equids, especially the common species
Beliajevina tekkayai. This is an animal with hypsodont teeth and thick coronal
cement, indicating a relatively wear-inducing diet, but preliminary study of its wear
facets shows that it was not exclusively a grazing animal (Fortelius, pers. comm.).
The second species of rhino is attributed to the genus Brachypotherium, but this is
represented by only one maxillary fragment, a mandibular tooth row, and a small
number of teeth, many of them broken. Its brachydont teeth indicate probable
browsing adaptation. A third species is attributed to Aceratherium (sensu lato) and is
represented by a few milk teeth.
Previous faunal lists for Pa~alar have included tapirs at the site, but this is
almost certainly based on a misidentified Listriodon molar (Hiinermann, personal
communication to Fortelius, 1988). We have no record of tapirs from our collection.
There is a very high level of breakage of the teeth of all the perissodactyl species
present at Patalar. It is rare to find complete Anchitherium or Brachypotherium
molars, and this is probably because they have little solidity. The high degree of
breakage of the rhino teeth is more surprising, although the more compact lower
molars which resist breakage better are more abundant than uppers. We have dis-
covered two rhino maxillae and several mandibles in the course of our excavations,
and in one of the gravel lenses there were the remains of both sides of a single
mandible, the bone of which had decayed in place leaving the tooth rows with crowns
intact but their roots missing. The common rhino Beliajevina is represented by a high
proportion of deciduous teeth, outnumbering permanent teeth by 31 to 7, but the
significance of this is uncertain. There is no evidence for carnivore activity directly
at the site; on the contrary there is every indication that carnivores were not primary
accumulators of bone at the site, but it is possible that the cachement areas from
which the bones were derived on the hillslopes above the site included a carnivove den
of some kind, and that the carnivore species preferentially collected juvenile rhinos at
450
its den. This is, of course, pure speculation, and there is no evidence on the bones
themselves of carnivore activity such as gnaw marks. The rhinos are under study by
Fortelius, and the equids by Forsten •
.Ariiodactyla
This is the most diverse group of mammals at P<tfalar, and some of its species
are by far the most abundant. Foremost of these is the small bovid Caprotragoides
which is well represented by horn cores, upper and lower jaws, teeth, and postcranial
bones. The morphology of the horn cores is rather primitive relative to those from
Fort Ternan and the Siwaliks, and the teeth in particular are primitive relative to the
Fort Ternan ones (Gentry, 1970). One other bovid species has so far been identified as
belonging to the genus Kubanotragus, a large species with relatively more hypsodont
teeth. Its horn core morphology is similar to that of Hypsodontus serbicus from
Prebeza (Pavlovic, 1969), and its dental morphology is specialized and similar to the
poorly known "Pachytragus" ligabue from the Hofuf Formation in Saudi Arabia
(Thomas, 1983). A number of horn cores and one partial skull are known for this
species, but only one mandible has been found. The teeth are very commonly broken,
more so than is seen for other bovids and giraffids. It can certainly be said to be
indicative of more open conditions at Pat~alar, but the primitive nature of the teeth of
Caprotragoides suggest that this animal lived in thicker vegetation than has so far
been suggested for Fort Ternan (Andrews et al., 1979; Shipman, 1986).
The giraffid Giraffokeryx is as abundant as Kubanotragus although neither

reaches the abundance of Caprotragoides. Formerly put into Palaeotragus, it is most
similar to members of the genus Giraffokeryx from Prebeza and Chios (Ciric and
Thenius, 1959). It is represented by several mandibles and an abundance of isolated
teeth, together with a few postcranial remains. Four other ruminant artiodactyls are
also present at Pafalar, represented only by isolated teeth. They. are the tragulid
Dorcatherium, a medium sized species the size of D. crassum. The early cervoid
Palaeomeryx is represented by just four teeth, similar to and slightly smaller than the
species from Sansan (Ginsburg, 1985). Finally, there are two cervid taxa, Micromeryx
and Euprox, both small with primitive browsing teeth. A more detailed study of the
ruminants is in preparation by Gentry.
Suoids form the next most abundant group after the bovids and giraffids. There
are two species of Listriodon, one bunodont and one lophodont, both common, although
only marginally more common than the hominoids. Identifications have so far been
restricted to unworn permanent cheek teeth, for which there are 100+ specimens
known for each species, and there are many so-far-unattributed specimens as well.
One species has similarities with L. splendens and the second probably represents a
new species. There is a small bunodont species, Conohyus simorrense (Hiinermann,
1975), which is less common and known from a single mandible and 35 teeth, and a
peccary attributed to Taucanamo. A few mandibles have been found of Listriodon and
Conohyus, but, as for the other taxa, most remains consist of isolated teeth. A more
detailed study is in preparation by Fortelius and Bernor.
Others
The fossil aardvaark Orycteropus is represented by a few teeth at Patalar. We

have also found two specimens of hyracoid, an isolated upper molar and incisor, and
these are considerably smaller than Pliohyrax graecus from Samos to which
Hiin~rmann (1985) tentatively assigned all Anatolian hyracoids. It is similar in general
size to Parapliohyrax mirabilis from Beni Mella! (Ginsburg, 1977).
TAPHONOMY
From the foregoing descriptions, it should be apparent that the fossil bones
recovered from the unsorted gravelly sands at PClfalar are extremely fragmentary.
Isolated teeth are the most commonly occurring skeletal part, and many of these are
451
broken as well. Similar degrees of breakage are present from the smallest to the
largest animals, with the sizes ranging from tiny insectivores to proboscideans. All
have similar proportions of isolated teeth to more complete specimens, and of broken
to complete teeth.
Bones in Relation to Sediment
Fossil bones are dispersed throughout the unsorted sands. Small mammal con-
centrations are highest in the middle to top of the unit, associated with fine sands
mixed with gravel; what we have designated as gravelly sand. Fine sand, silt, and clay
make up at least 50% of these deposits, and gravel and pebbles make up 6-14%. Large
mammals are rare, but are abundant in what we have called coarse sands where small
mammals are almost absent. These consist of 45-50% coarse sand and only 30-35%
finer elements. These are the two most abundant sediment types, although clearly
they only differ in degree. More different still are the gravel lenses, and these are
clearly definable in the field. They consist of up to ZO% gravel and pebbles, 50%
coarse sand, and less than 30% finer fractions. They may be rich in fossils or com-
pletely unfossiliferous, but only large mammal bones are found, some of them rela-
tively complete (for example, the partial bovid skull (figure 3), a proboscidean skull,
an Amphicyon mandible, and the complete rhino tooth rows, all mentioned earlier).
Bone orientations are random in the gravel and fine sands, but have preferred
directions in the coarse sands. Where the coarse sands surround a gravel pipe, the
orientations of the bones in the coarse sand run parallel to the edges of the gravel
pipe, and where the coarse sand is sandwiched between two gravel lenses, the bones
are oriented longitudinally between the edges of the lenses. Most of the bones have
low angles of dip except in the gravel pipes, where there is a high frequency of near-
vertical bones. This disposition of the bones in the sediment by size and orientation
shows that the bones accumulated at Pa.!jalar as a direct result of sedimentary activ-
ity. Bone distribution is thus a secondary feature, relating to physical processes of
sediment accumulation rather than to any direct biological activity.
Pre-Depositional Evidence
No direct evidence of carnivore activity has been observed on any of the larger
bones in the unsorted sands at Pa!falar. Indirect evidence may be seen in the 4 to 1
preponderance of immature individuals of the rhino Beliajevina that has been com-
mented on earlier, and it is hard to explain this, other than as the result of carnivore
selection. For the small mammals, a low percentage of rodent teeth has earlier been
observed to have been affected by digestion, and these must have derived from pellets
or scats of a Category 1 predator living within the cachement area for the site. There
is insufficient evidence as yet to identify the predator (Andrews, 1989), although it is
clearly neither a diurnal raptor nor one of the carnivores that produce extreme diges-
tion (canids and felids).
Many of the bones show signs of weathering on their surfaces, and some have
traces of root marks. In both cases, the bones affected are dispersed throughout the
deposit, confirming that the alteration did not take place after deposition. There is a
high correlation between weathered bone and evidence of rounding. It has been
described earlier how a moderate percentage of bone from all species shows evidence
of attrition producing rounded surfaces, but all of this bone can be seen to have been
weathered originally. In some simulation experiments on recent bone conducted with
different grades of Pafalar sediment, we have found that weathered bone becomes
rounded by attrition very much faster than does fresh bone; the more heavily
weathered the bone, the m~re rapid the rounding, and the effects of the attrition are
such as to obscure the weathering, since all the loose and flaking bone which charac-
terizes weathered bone is worn away. Some rounding of weathered bone has been
found to occur within a very short period, so that the degree of rounding present on
most of the bones found at P~alar does not indicate prolonged water action, as has
formerly been thought, but is consistent with short term attrition of weathered bone.
Some bones, however, do show evidence of prolonged attrition, and these have been
preserved in stream channels within the cachement area of the Pa,alar sediments.
452
Fig. 3. Top, an accumulation of large bones in one of the gravelly
lenses associated with potholes, showing part of the skull of
Kubanotragus and a broken mandible of Amphicyon; below,
an oblique view into the excavated bottom of the pothole
illustrated in figure Z in which most of the dentition and
parts of the skull of a single individual of Gomphotherium
were found.
The presence of weathering itself is consistent with the model of sediment

accumulation described earlier. It is inferred that bone was accumulating on the
surface of the hillslopes and in streambeds above but close to the present site of
Pa~alar, and depending on the age of the bone all stages of weathering would be repre-
sented together with some rolled bone . Most bone had at least some degree of
weathering, which would have weakened it in proportion to degree, and probably there
was already a high proportion of isolated teeth left after destruction of jaws by
weathering. Broken tooth fragments frequently show evidence of weathering, most
apparent on the proboscidean teeth, so that much of the breakage can be shown to
have been pre-depositionar. The transport of the bone during the flooding that pro-
453
duced the accumulation of P~alar sediments would have completed the destruction of
most of the bone and produced the rounding evident on the more weathered bone,
leaving the teeth and fresh bone relatively intact. Any bone present on the ground
would have been affected in this way, so that there was no selection of bone size or
species other than by the depositional processes described in the preceding section.
Only the strongest elements such as teeth, mandibles, horn cores, and some post-
cranial elements have survived these processes to form the bone accumulation as we
see it today. The bone assemblage was thus effectively a death assemblage represent-
ing animals that died on the hillslopes overlooking the site, with the addition of an
unknown but probably small proportion introduced by predators.
PALEOECOLOGY
In attempting to define the paleoecology of the Pasalar fauna, we have built up a
wide range of comparative faunas with which to compare it. These include the 23
recent tropical African faunas described by Andrews et al. (1979) together with 28
additional faunas from Europe and Asia. These include several from Turkey itself,
covering forest and steppe conditions, but there is clearly no habitat in Turkey today
that is directly comparable with conditions in the Miocene. We therefore extended
our range of investigation to cover woodland/forest types all over Eurasia, concen-
trating on the richer deciduous forests of central and southern Europe. Some more
impoverished coniferous forests were also investigated, including the northern boreal
forests, and some tundra habitats were also examined. Finally, the sample was
extended to cover the subtropical monsoon forests of India, semi-deciduous and with a
definite dry season, and the tropical forests of southwest China. The original data for
this comparative sample will be presented elsewhere, but the information can be
summarized here for comparison with the Pasalar fauna.
The basis for the comparison of fossil with recent faunas is the analysis of
ecological diversity (Andrews et al., 1979). This relates the range of adaptations
present in mammalian.communities to habitat types on an empirical basis. The adap-
tive strategies of mammals are assessed in terms of body size, diet and locomotion,
and the differing proportions of these strategies in communities from differing
habitats are used to categorize and distinguish the habitat types. The analysis is based
on numbers of species because comparative information on numbers of individuals of
each species is not available. The total numbers of species is a significant factor in
the analysis, although by itself it is only a crude measure of diversity. Also useful in
some contexts is the taxonomic composition of the faunas. Consistent patterns of
species diversity and adaptive strategy are found for mammalian communities derived
from similar habitats, and this consistency has been attributed to stability of trophic
guilds through time (Olson, 1966) and across geographically disjunct regions (Harrison,
1962; Andrews et al., 1979). The patterns of interaction in mammalian communities
are more distinctive and longer lasting than the ephemeral presence/absence of indi-
vidual species, and so the structure of the communities provides a better guide to the
nature of the fauna and its ecology. Where many or all of the species are extinct, as
is the case for fossil faunas, individual species preferences cannot be known in any
case, so that the analysis of community structure of the fossil faunas offers an alter-
native approach.
The Piifilar Fauna
Previous interpretations of Pa~alar paleoecology have indicated dry woodland to

steppe conditions during the Miocene (Becker-Platen et al., 197 5). Schmidt-Kittler
(1976) essentially agreed with this interpretation, stating that the environment may
have been even more steppe-like on the basis of hyaenids in the fauna, but he had
difficulty reconciling this view with the high diversity of canoid (= Arctoidea of
Schmidt-Kittler) carnivores; 9 out of 15 species belong to this group, and this is more
indicative of closed forest conditions (Schmidt-Kittler, 1976). Our analysis is based on
the 53 species recorded in Table 1, and the first observation to be made on this is that
it is a high level of species diversity compared with present-day temperate faunas,
454
much greater than is found in the richest forest faunas. It is also higher than is found
in tropical savannas and grassland (see below), and this of itself is sufficient to cast
doubt on the previous interpretations of paleoecology.
The ecological diversity of community structure of the Papalar fauna is shown

graphically in figure 4. The Pilfalar fauna is shown compared with means of faunas
from four recent habitat types, as follows:
1. Semi-deciduous, summer-rainfall seasonal forest in sub-tropical India, often called

monsoon forest, with a single main tree canopy dominated by one tree species, a
discontinuous lower tree canopy, and rich ground vegetation, with the trees inter-
spersed with extensive herb- and grass-rich meadows.
z. Deciduous forest, highly seasonal with a long dry season in tropical Africa, usually
referred to as tropical woodland; there is a single tree canopy, dominated by one or
a few species, no lower canopy but grass- and shrub-rich ground vegetation.
3. Deciduous temperate forest, with a wet winter and hot dry summer in southern
Europe, often mixed altitudinally with broad-leaved trees at lower altitudes and
coniferous trees at higher altitudes; there is a single tree canopy and herb-rich
ground vegetation.
4. Semi-deciduous tropical rain forest in Africa, seasonal, with a moderately long dry
season; there are two to three tree canopies, with moderate species diversity,
greater than any of the preceding examples, but much less than the tree species
diversity of wetter evergreen rain forest types.
The African data come from Andrews et al. (1979) and the Indian and European
data from material assembled by C. Artemiou (1984). Four analyses are shown,
taxonomic, size, locomotor, and dietary. The taxonomic analysis simply shows the
overall taxonomic compatibility of the Pa9alar fauna with those living today, and it
supports the taphonomic evidence that the Pilfalar fauna is not heavily biased with
respect to any one taxonomic group.
The analysis of body size shows a high level of equitability in the Pi!falar
mammalian fauna (figure 4). The body size classes are more evenly filled than are
those of any of the recent faunas, which makes it hard to interpret the significance of
this feature. It may be observed that in comparison with the recent faunas, those
from tropical to subtropical seasonal forests have the highest levels of equitability, so
that the Payalar pattern is closest to these, but there are evident differences in size
structure that cannot be assessed at present. It appears that there is a size bias
against the smallest rodent and insectivore species, which has already been seen to
have been indicated by the taphonomic evidence, but it is possible that when the work
on the small mammals has been completed the frequency of class A species (0-100 g)
may increase.
The locomotor structure of the fauna is dominated by terrestrial species (figure

4). Large ground mammals, by which is meant larger species that spend all of their
time on the ground, constitute more than 50% of the mammalian species. Another
30% of the fauna consists of small ground mammals, and this encompasses the smaller
mammals that habitually live on the surface of the ground, but it also includes those
species living in shallow holes (excluding fossorial species), in banks, rocky areas, low
bushes, fallen trees, and so on. Together these terrestrial species make up 80% of the
fauna from Payalar. Arboreal and scansorial species are present in low but significant
numbers, but other classes of species, namely those with aquatic, fossorial, and aerial
adaptations, are poorly represented.
The dietary structure of the Pilfalar fauna shows a high degree of equitability, in
contrast to the locomotor structure just described (figure 4). Carnivorous species are
most abundant, as might be expected from the high representation of Carnivora in the
fauna. Browsing herbivores and frugivores are the next most abundant species types,
455
TAXONOMIC SIZE LOCOMOTOR FEEDING
ORDER CATEGORY ADAPTATION ADAPTATION
%
so
40
PAS ALAR
30
20
10
so
40
Subtropical semi-
lt
30
deciduous forest
20
N=2 10
50
40
Li
Traplcal deciduous
30
IL
forest
20
N=3 10
u
50
40
Temperate mixed
30
forest
20
N=7 10
[~La
Traplcal semi-
lb
evergreen forest
N•3
R I PACO IFBGCO
Fig. 4. Community structure of mammalian faunas, showing the means from four
recent habitat types compared with that of the Pllfalar mammalian fauna.
The recent faunas represent, from bottom to top, the mean of 3 semi-
evergreen tropical rain forest environments in tropical Africa (Andrews et
al., 1979), the mean of 7 deciduous and mixed forests from southern
temperate regions of Turkey, southern Germany, southern France and Spain,
the mean of 3 deciduous forest/woodland environments in tropical Africa,
and the mean of Z semi-deciduous forest areas in the sub-tropical zone of
India. These habitats are described more fully in the text. The vertical
scale shows the percentage numbers of species. The four categories are as
follows: Taxonomic, R, Rodentia, I, Jnsectivora, P, Primates, A,
Artiodactyla, C, Carnivora, and O, others; Size, A, 0-100 g, B, 100 g- 1 kg,
C, 1-10 kg, D, 10-45 kg, E, 45-180 kg, and F, >180 kg; Locomotor, L, larger
mammals wholly confined to the ground, S, smaller mammals that are
mainly terrestrial but also live on the lower parts of trees and bushes and on
fallen trees, A, arboreal, Sc, scansorial, and O, others (fossorial, aerial,
aquatic); Feeding, I, insectivorous, F, frugivorous, B, browsing herbivory, G,
grazing herbivory, C, carnivorous, and 0 1 omnivorous.
456
with grazing herbivores considerably less commonly represented than browsers. This
proportion, and the relatively high proportions of frugivores, are the two significant
aspects of the dietary adaptations of the Pafalar fauna.
Comparison with Modern Communities
Looking at these modern communities in more detail now, their differences will
be documented in terms of the adaptive strategies mentioned earlier. The most
obvious difference is to be seen in the total numbers of species present in the commu-
nities. The 53 species present in the P~alar fauna place it well beyond the limits of
most temperate faunas but short of most tropical faunas. It is most similar, assuming
no species loss, to subtropical forest faunas from India.
The taxonomic composition of the Pafalar fauna is most similar to that seen
today in tropical woodlands, since they share the high numbers of carnivores, fewer
rodents, and very few primates (figure 4). The P~alar fauna also bas a high level of
size equitability, although the pattern is not particularly close to any of the recent
faunas. This is because of the lower proportion of mammals in the 1-10 kg size range,
and this might suggest the presence of a bias in the size composition of the Patalar
fauna. This apart, it may be said to compare most closely with the size structure of
tropical deciduous or subtropical semi-deciduous forest faunas (figure 4).
The Pa~alar fauna clearly differs from the tropical evergreen forest pattern in
its locomotor diversity, because it has 79% terrestrial species dominated by large
mammals wholly restricted to ground living. In this also i.t is most similar to the
tropical deciduous or subtropical semi-deciduous forest faunas (figure 4). Differences
in dietary adaptations provide even more marked contrasts between mammalian
communities and provide the best discrimination between faunas from different
habitat types. Discrimination is even possible between the less rich tropical forest
faunas, and deciduous single-canopy forest faunas, with the subtropical summer-
rainfall forest faunas falling at an intermediate position as a result of increases in
proportions of browsers and grazers relative to tropical forest faunas, and increases in
frugivores and decreases in grazers relative to deciduous forest faunas. The Patalar
fauna dietary structure is particularly close to that of the monsoon summer-rainfall
forests with its intermediate levels of frugivorous species but lower proportions of
grazers (figure 4).
This comparative series shows very clearly the affinities of the Patalar fauna.
Its community structure may be distinguished from that typical of both temperate and
tropical forests in all the analyses. It also differs widely from steppe, semi-desert,
and other open country community structures. Consistent similarities are found with
faunas from both tropical deciduous forest/woodland and subtropical semi-deciduous
forest in the summer rainfall belt, and more with the latter than with the former.
These habitats have a certain amount in common, and it may be inferred from this
consistency of pattern that the climate at Pafalar during the Miocene was subtropical
at the very least, with a pronounced dry season. Less certainly it may be inferred that
the rainfall pattern was one of summer rainfall, i.e., monsoon conditions, since the
present climatic regime of winter rainfall in Mediterranean regions supports very
different community structures of the mammalian faunas. The vegetation supported
by this climate was probably similar to the Sal forests of northern central India today,
closed forest, single species dominated, deciduous to semi-deciduous, interspersed
with grass meadows. This evidence is in accord with evidence from the pollen record
for this part of the middle Miocene (Benda, 1971).
ACKNOWLEDGMENTS
We are very grateful to the many people who have helped in the fieldwork at
P8.falar. This includes particularly Ayban Ersoy who has worked on aspects of the
taphonomy, Mikael Fortelius, who excavated the pothole illustrated in figure 3 and bas
provided the notes on the rhinos and pigs, and Alan Gentry and Louis Jacobs, who have
457
provided information on the ruminants and small mammals respectively. We are also
grateful to the organizers of the NATO workshop for their invitation to attend and for
their excellent organization of the meeting.
The Palfalar excavation has been funded by the Directorate of Antiquities, T.C.
Ministry of Culture and Tourism. Financial support for the participation of non-
Turkish personnel has been consistently and generously provided by the Leakey
Foundation, with assistance for specific projects from the Boise Fund. The Pafalar
excavation has been carried out on behalf of the T. C. Ministry of Culture and
Tourism, Directorate of Antiquities, and the University of Ankara, Faculty of
Language, History and Geography, Department of Palaeoanthropology.
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459
TAPHONOMIC AND SEDIMENTARY FAC'l'ORSIN THE
FOSSU. RECORD OF MAMMALS
M. A. Alvarez Sierra1 , M. Diaz Molina1,

J. L Lacombaz, and N. L6pez Martmez1
1Departamento de Paleontologfa
UEI de Paleontologfa
Instituto de Geologfa Economica del CSIC
Facultad de Ciencias Geologicas
Universidad Complutense
Z8040 Madrid, Spain
ZDepartamento de Geologla
Facultad de Ciencias Biologicas
Universidad de Valencia
Burjasot, Valencia Spain
INTRODUC"'ION
The composition of the fossil assemblages of mammals (MFA) is traditionally

considered as being controlled by historical factors. Its variations are attributed to
biochronological changes due to evolutionary process or to biogeographic changes due
to geodynamic process. The different MFA are compared as sampling units of similar
nature. The presence or absence of taxa and their relative abundance are used as
biochronologic and biogeographic indicators, and the samples (MFA) are considered
homologous, although they appear in a different sedimentological, taphonomical, or
tectonic context.
More recently, processes other than historical ones have been considered as a
source of variability of the MFA. That is the case of the paleoecological and paleo-
climatic factors, mainly changes in humidity and temperature. Several authors (van
de Weerd et al., 1978; Daams et al., 1984; Hugueney, 1984) have shown that the MFA
composition depends on these factors in many cases. Different sites of the same age
(e.g., karstic and basinal sites) may show different composition, because paleoecolog-
ical conditions (e.g., aridity) differ in both types of landscapes.
Tectonics seem also related to the variability of the MFA composition, since
many MFA are included in diastrophic basins with frequent changes in the geographic
and sedimentary pattern. Tectonic events are claimed to be the main factors control-
ling the composition of MFA according to De~ys et al. ,(1985, 1986); on the contrary,
they seem not directly related according to Lopez Martinez et al. (1985, 1987), tecto-
sedimentary breaks occur often in the middle of the biozones. Nevertheless, diastro-
phism causes frequent autophagic processes, reworking sediments and fossils, and it
may affect also the MFA. That has rarely been recognized by vertebrate paleontol-
ogists, but is a common problem for invertebrate and plant fossil assemblages.

We have approached in this note the study of the taphonomic factors affecting
the homogeneity of the MFA. Taphonomic factors are the latest in the formative
processes of the MFA, and therefore they may be examined first. Tectonic and paleo-
climatic factors act in terms of large-scale processes affecting the MFA formation
via the sedimentary and taphonomic processes. We study these factors for fossil
assemblages of small mammals (mMFA) which are far more abundant, but their results
may be useful also for the study of the large mammal fossil assemblages (!MFA).
METHODS
Three stratigraphic sections have yielded the mMFAs used in our study. One is
situated in the northeastern Ebro basin (Alvarez Sierra et al., in press); it has some
1000 m thickness and contains three mMFAs extending from the Oligocene-Miocene
boundary interval to the early Miocene (see Alvarez Sierra et al., 1987). The second
section is situated in the western Ebro basin (Cuenca, 1985; Martinez, 1987), yielding
five mMFAs from the late Oligocene to the early Miocene, with about 300m thick-
ness. The last section is situated in the Loranca basin (Pfaz et al., 1985; Daams et al.,
1986) and contains seven mMFAs from the late Oligocene to the early Miocene in
some 600 m thickness. Tecto-sedimentary breaks are intercalated in the three
sections, in the Oligo-Miocene boundary interval and in the early Miocene (Lopez
Martinez et al., 1987). The systematic and biochronologic study of the rodents has
been done by Cuenca (1985), Martinez (1987), Alvarez Sierra (1987), and Lacomba
(1988).
The localities have been classified according to the sedimentary facies and
stratigraphic sequences containing them. Five types of environments have been
interpreted from these data. They have been represented in figure 1, and may be
defined as follows:
1. Channels. They are inferred from the sedimentary bodies with an erosional basal
surface formed by stream incision. Grain size varies from gravel to silt.
z. Floodbasin. They are the lowest lying parts of a stream flood plain. Suspended
fine sediments settle from overbank flows. Grain size varies between silt and silty
clay.
3. Ponds. They are relatively small, episodic bodies of fresh or brackish stagnant
waters. They are interpreted from thin and discontinuous calcareous deposits
associated with flood plain sediments or oxbox-lake fill.
4. Swamp-Marsh. More permanent, anoxic stagnant waters with a rich plant cover
(trees or grass). They are recognized from the peat content of the sediment.
5. Lakes. They are large, permanent water bodies. They are distinguished on the
basis of thick and extensive calcareous deposits.
Table 1 shows the distribution of the 15 main localities in these five types of
environments. We have included two other mMFAs, Cetina and Bunol, which are
isolated in two different small basins. They are well known in the paleontological
literature (see Daams, 1976) and a taphonomical study of the Builollarge mammals has
been done by Belinc:fton (1987).
The preservation of the micromammal remains has been studied on the isolated
teeth. Much relevant information comes from the bones (Behrensmeyer, 1978; Korth,
1979), but they are rare in the basinal mMFA and determinations are difficult. The
procedure of the taphonomic study of small mammal teeth is from Lacomba (1988).
Five preservation states are distinguished:
AI Molars with complete crowns (with or without roots) and with intact enamel.
462
STRATIGRAPHIC
ENVIRONMENT SUCCESSION SEDIMENTS LEGEND
FOSSIL CONTENT
§}
silt to silty 0 Micromammals
clay
~ Gastropods
. ~ Charophytes
,..... Ostracodes
! -~~·'/1
thin and
discontinuous l!Yl Plant remains
calcareous
~j
& deposits
LITHOLOGY
0 dl• ....... channel-fill
Coarse terrigenous
D sediments
. ./ ,_;J .,~' lake thick and
extensive c==J
IT::]
__, .. · .,<·/'. ).. ·. ., ~ calcareous Silt and silty clay
Marl
1 depooHo tt"2J
.,,
~~-
~"1" 1'·,,,, \1.
I O.t•~<~ - Limestone
Peaty clay
swamp-marsh ~ dJB~
~H~ peaty clay OTHERS
Q ~..#-.:;,"1"'1-,~o. ""-,~-~....
- ~Crevasse-splay
c,_~~~~-1~....:-....~ ~ o~!J~
~~~~~-----4--------~---------t ~-~Point bar deposits
"'-=-~-!Clay plug (oxbow deposits)

sand C Ox bow lake , e Oncoli tes
~0 [:~~:-;~ Palaeochannel,~ Roots
.,.
CJ)
Fig. 1. Classification of the different sedimentary environments, and their stratigraphic successions .
Co)
Table 1. Distribution of the studied localities according to its age, procedence and
type of environment. Olig.-Mioc. b.i. means Oligocene-Miocene boundary
interval (see Alvarez Sierra et al., 1987). Kg is the kg of sediments treated
for each locality.
Loc. Environment
Number Loc. Name Age Zone Basin Type Kg
1 PAR Parr ales late Oligocene u Loranca Flood basin 1.000

Z BG Bergasa late Oligocene w WEbro Pond Z.lZS
3 MON Moncalvillo Olig.-Mioc. b.i. X Loranca Pond 6.000
4 AU Autol Olig.-Mioc. b.i. X WEbro Lake 550
5 sc Santa Cilia Olig.-Mioc. b.i. X NE Ebro Pond 3.1Z5
6QU1 Quell Olig.-Mioc. b.i. yl WEbro Lake 1.575
7 PZ Pozo Olig.-Mioc. b.i. Yt Loranca Pond szs
8 CE Cetina Olig.-Mioc. b.i. Yt Almazan Swamp 50
9 FM Fuenmayor Olig.-Mioc. b.i. yl W Ebro Pond l.ZSO
10 MOH Moheda Olig.-Mioc. b.i. Yt Loranca Pond 3.ZOO
11 VQ Valquemado Olig.-Mioc. b.i. Yt Loranca Swamp 700
lZ CAR Carre til early Miocene Yz WEbro Lake 375
13 AC Alcocer early Miocene Yz Loranca Pond 1.7Z5
14 SJ San Juan early Miocene Yz NW Ebro Channel 900
15 GAL La Galocha 5 early Miocene z NW Ebro Pond 750
16 HO Huerta Obispalia early Miocene z Loranca Swamp z.soo
17 BU Buiiol early Miocene c Buiiol Swamp 500
BII Molars with fracture or corrosion traces affecting the enamel of the crown.
There are no traces of rounding.
CII Molars with fracture or corrosion traces affecting the dentine of the crown.
There are no traces of rounding.
Bill Molars with damaged crowns presenting rounded erosional surfaces affecting the
enamel.
em Molars with damaged crowns presenting rounded erosional surfaces affecting the
dentine.
For the statistical treatment of the data between the five types of preservation
defined, these have been grouped into the following three preservation classes:
I Molars with intact crowns (enamel and also dentine).
II Molars with damaged enamel or dentine of the crown but without traces of
rounded erosional surfaces (BII plus ZCII).
m Molars with damaged enamel or dentine of the crown presenting traces of

rounded erosional surfaces (Bm plus ZCIII).
For each assemblage we have examined the preservation state of the· first and
second molars of two rodent families, the Gliridae and the Eomyidae. Both have
brachyodont to semihypsodont crowns of small to medium size (around 1 to 3 mm).
Their mechanical properties are therefore considered to be similar.
The analysis of data gives an idea of the relation between the sedimentary
environment and the preservation of the micromammal remains, with a view to evalu-
ating the homogeneity of the conditions of the mMF A formation. A more extensive
464
study will be necessary to conclude the main factors controlling the composition of
the mMF A. However, some interesting remarks may be extracted from this prelim-
inary analysis.
TAPHONO~CFACTORS
Taphonomy means "the laws of burial," and was originally described as the study
of the transition of the organic remains from the biosphere into the lithosphere
(Efremov, 1940). That field has been comparatively more developed for invertebrate
than for vertebrate paleontology, but it is becoming a current topic in many research
programs on vertebrate studies.
Fossilization process may be divided by three events: production, burial, and

record. Taphonomy distinguishes usually among biostratinomy, study of the fossiliza-
tion processes before burial, and fossil diageneis after burial (Lawrence, 1971). The
production of micromammal remains may be due to predatory activity (as fecal or
rejection pellets) and also to attritional (diseases, senility) and mass mortality (catas-
trophes as starvation, pests, floods, etc.). Many agents can modify the state of the
remains and may drive often the remains at a place different than that of the produc-
tion one (nekrokynesis). The remains sedimented after the production are called
accumulated. The remains may be removed one or more times during the biostrati-
nomic phase, and then they are considered redeposited. After burial, the remains
which are removed during the fossil diagenetic phase are considered reworked
(Fernandez L-6pez, 1981, 1987).
Reworked remains have been rarely mentioned in the vertebrate paleontology

literature. Micromammal remains are unlikely to be reworked after fossil diagenesis
because of their fragility, but reworking has been observed in other microfossils and,
therefore, this contingency seems .! priori not impossible. The frequence of auto-
phagic sedimentation in syntectonic filling basins increase the possibility of reworked
remains being mixed with accumulated-resedimented ones; even complete assemblages
could be reworked and buried in younger sediments.
Since microfossils contain little or no matrix sediment in their cavities, the main
criteria to distinguish between accumulated, resedimented, and reworked sediments in
mMFA will be the conditions of preservation. We will complete our inferences with
other criteria, as the upper/lower molars ratio (Voorhies, 1969), and the study of the
intraspecific variability.
RESULTS
Tables Z and 3 show the observed frequencies of the glirid and eomyid molars for
the different preservation states. For each table, the upper part shows the number of
specimens, and the lower part the percentages for each locality. The total number of
specimens is similar for both families (963 glirids and 958 eomyids), and the preserva-
tion is generally the same (454/401 intact, Z94/3Z7 in Bll, 107 both in en, 84/81 in
Bm, and Z4/4Z in em). However, there are different and even contradictory distribu-
tions of the preservation states of the families in each locality. That is an unexpected
result, which will be discussed later because it does not fit with the commonly
accepted hypothesis that most micromammal remains are produced by predatory
activity (Mellet, 1974; Mayhew, 1977; Dodson et al., 1979; Andrews and Evans, 1983).
The comparisons among the different localities has been done in figures Z and
3. Each locality is placed in a scatter diagram according to its value of the ll and m
preservation classes. The class ll represented in the Y-axis may be related to altera-
tions or breakage produced by chemical or mechanical actions (gastric or humic acids,
roots growing, impact, etc.). The class m represented in the X-axis may be related to
mechanical erosion mainly attributed to transport by tractive currents. The class I
(intact elements) is not shown, being represented by the relative distance of each
465
Table Z. Frequencies of the glirid molars for the different preservation states. The
upper part shows the numbers of specimens, and the lower part the percent-
ages for each locality. Loc. number is locality number, the same as in table
1.
Loc.
Number AI BIT en Bm em Total
1 PAR 4 4 3 3 3 17
Z BG 63 Z6 6 7 0 10Z
3 MON Z8 30 1Z 1 0 71
4 AU 31 15 z 4 0 5Z
5 sc 38 Z7 5 1 0 71
6QU1 Z8 14 z z 0 46
7 PZ 5 6 3 4 z zo
8 CE 14 4 1 1 0 zo
9 FM 6 3 z 3 z 16
10 MOH 4Z 45 11 z 0 100
11 VQ Z9 Z9 9 14 z 83
1Z CAR 1Z 7 z 1 0 zz
13 AC 17 7 3 3 0 30
14 SJ 30 18 18 13 5 84
15 GAL 30 18 9 8 4 69
16 HO 18 Z5 8 6 z 59
17 BU 59 16 11 11 4 101
Total 454 Z94 107 84 Z4 963
Loc.
Number AI BII err Bm em Total
1 PAR Z6 Z6 zo 14 14 100
Z BG 6Z Z5 6 7 0 100
3 MON 40 4Z 17 1 0 100
4 AU 60 Z9 7 4 0 100
5 sc 53 38 1 8 0 100
6QU1 60 30 5 5 0 100
7 PZ Z5 30 15 zo 10 100
8 CE 70 zo 5 5 0 100
9 FM 38 19 1Z 19 1Z 100
10 MOH 4Z 45 11 z 0 100
11 VQ 35 35 17 10 3 100
1Z CAR 55 3Z 8 5 0 100
13 AC 56 Z4 10 10 0 100
14 SJ 35 Z1 15 zz 7 100
15 GAL 43 Z6 13 1Z 6 100
16 HO 31 4Z 13 10 4 100
17 BU 58 16 11 11 4 100
Total 789 500 186 165 60 1700
466
Table 3. Frequencies of the eomyid molars for the different preservation states. The
upper part shows the numbers of specimens and the lower part the percent-
ages for each locality. Loc. number is locality number, the same as in table
1.
Loc.
Number AI Brr err sm em Total
1 PAR 16 1Z z 1 z 33
Z BG Z4 17 6 3 1 51
3 MON 17 30 9 11 11 78
4 AU 40 44 Z6 15 10 135
5 sc 40 35 7 11 7 100
6 QU1 31 19 10 10 z n
7 PZ 4 10 4 0 0 18
8 CE 5Z 19 3 4 0 78
9 FM 8 3 0 0 0 11
10 MOH 7 1Z 3 4 z Z8
11 VQ 5 7 4 0 0 16
1Z CAR Zl zo 6 3 3 53
13 AC 69 39 10 3 1 1ZZ
14 SJ 3 8 0 0 0 11
15 GAL z z 1 z 0 6
16 HO zo 13 9 3 1 46
17 BU 4Z 37 7 1Z z 100
Total 401 3Z7 107 81 4Z 958
Loc.
Number AI Brr err sm em Total
1 PAR 49 36 6 3 6 100
Z BG 47 33 lZ 6 z 100
3 MON zz 38 lZ 14 14 100
4 AU 30 33 19 11 7 100
5 sc 40 35 7 11 7 100
6QU1 43 Z6 14 14 3 100
7 PZ zz 56 zz 0 0 100
8 CE 67 Z4 4 5 0 100
9 FM 73 Z7 0 9 0 100
10 MOH Z5 43 11 14 7 100
11 VQ 31 44 Z5 0 0 100
lZ CAR 40 38 11 6 5 100
13 AC 57 3Z 8 z 1 100
14 SJ Z7 73 0 0 0 100
15 GAL 33 33 17 17 0 100
16 HO 43 Z8 zo 7 z 100
17 BU 4Z 37 7 1Z z 100
Total 691 636 195 1ZZ 56 1700
467
GLIRIDAE
80
CHANNEL 0
MON
FLOODBASIN !::.
0 POND 0
LAKE Jt
••
70 VAQHO
MOH PAR SWAMP-MARSH •
0
BG
PZ
60 0 0
CAN SJ
0 0
...
50 CAR
AU QC FM
...QIJ1 sc 0
BU
•
40 ... 0
H
H
g
•
+ CE
~ 30
"'
20
10
VII
10 15 20 25 30 35 40 45 50
~----·
200
% BIII+2 CIII
Fig. Z. Comparison of the degree of preservation of Gliridae associations from

the various localities. The vertical axis represents the percentage of
elements that present the stage of preservation II (BII plus ZCII), and in
the horizontal axis the percentage of elements that present the stage of
preservation m (Bm plus ZCIU). The horizontal axis differentiates three
locality-groups from still water environments to turbulent current
deposits.
468
1001 PZ
• VAQ
:~
EOMYIDAE
Channel D
AU Floodbasin 6.
OSJ
HO GAL * Pond 0
70
•
60
0
*CAR
o MOH Lake
Swamp-Marsh • *
........ BGO
BU *QU1 0 MON
0
50 AC
•
-
(\1
0 oSC
+ b. PAR
m 40
0:..!! VIL
• • CE
30bFM
20
10
0
5 10 15 20 25 30 35 40 45 50 55
% Bill t 2Cl1I
Fig. 3. Comparison of the degree of preservation of Eomyidae associations

from the various localities. The vertical axis represents the percent-
age of elements that present the state of preservation II (BII plus
zcm, and the horizontal axis the percentage of elements that
present the stage of preservation m (Bill plus ZCDI). In the hori-
zontal axis no such well-differentiated locality-groups are observed
as the case in the Gliridae.
locality to the coordinates origin. It may be related to accumulated, unaltered

elements.
Figure Z corresponds to the glirid assemblages. The X-axis differentiates the

localities into three groups. First group to the left contains the three lacustrine, one
swamp, and several pond-related sites; second group has most of the swamp sites; and
finally, the third group to the right has the fluviatile localities. Therefore, the m
preservation class seems related to the hydrodynamic energy of the sedimentation; the
highest values of rounded elements correspond to the flowing water sites. Conse-
quently, the observations strongly support the hypothesis that the hydrodynamic
currents cause the rounded erosion surfaces of the teeth.
Figure 3 corresponds to the eomyid assemblages. Groups of localities are less

well defined, and the general trend of the environments is contrary to that observed in
the case of the glirids.
The inverse ratio of the preservation state of glirids and eomyids assemblages
may be seen in figure 4. This diagram shows the values of the category m index of
eomyids (Y-axis) and glirids (X -axis) for each locality. A linear regression has been
469
I CAN
0
50
H MON
H
H
u 40
(\J
+
H
H
H 30
"
ill
MOH SC
w o.._ o
,.•
<I;
Q '-. AU
H
>-<
:;:: 20 QU1~'\.
GAL
•
0
w CAR'-. BU PAR
0
10
BG " ' '-. HO
D.
0
'\.
•
CE AC '\.
0
VAQ ' " ,...--- SJ0 0PZ 0
FM
0
10 20 ' '30 40
---------
50
GLIRIDAE
60 70
BIII + 2CIII
Fig. 4. Comparison between the preservation stage m (BID plus ZCIII) in the
different localities of Eomyidae (vertical axis) and Gliridae (horizontal
axis) associations. Note the inverse relation between the preservation
stage m (evidence of transport) of Gliridae and Eomyidae from the
same localities.
calculated to show the negative slope and the inverse relation between both varia-
bles. The correlation coefficient is r:-0.588, and it seems that it will be higher in the
case of a curve regression.
Two small samples coming from the Loranca basin, Villalba (swamp) and Canales
(pond), have been added to the diagram to show the clear phenomenon of the divergent
preservation state of the glirid and eomyid teeth. Even these non-t'epresentative
samples, with only 3 and 14 specimens each, shows opposite states of conservation in
both families.
The differences in the distribution of the preservation classes suggest that the
glirids and the eomyids were affected by different processes and did not arrive at the
locality in the same way. If their remains have been produced by predators, their
preservation states might be similar, since the physical features of their bones and
teeth are also similar. The majority of the studied assemblages contain both accumu-
lated and redeposited elements, except some localities as Cetina. These mixed type
of assemblages suggest that predation should not be invoked as the main mechanism in
the formation of basinal mMFA.
470
Table 4. Frequency pattern of upper and lower molars of Gliridae and Eomyidae
species.
Locality Species N of Ml-Z %of Ml-Z/
Gliridae
Bergasa Ebromys bergasensis 191 49
Moncalvillo Pseudodry"om}'S ibericus 30 50
Perid:yromys murinus 75 56
Santa Cilia Pseudodryom}'S ibericus llZ 46
Peridyromys murinus 67 53
Vasseurom}'S rugosus 6Z 53
Vasseuromys priscus 85 48
Quell Quercom}'S bijmai 59 54
Valquemado Pseudodryomyslopezae 103 47
Moheda Pseudodry"om}'S lopezae 140 53
Perid:yromys murinus Z8 50
Eomyidae
Bergasa Rhodanomys transiens 1137 45
Pseudotheridom}'S schaubi 34 44
Moncalvillo Rhodanomys schlosseri 343 48
Autol Rhodanomys schlosseri Z69 5Z
Santa Cilia Rhodanomys oscensis 908 5Z
Quell Ritteneria molinae 139 5Z
Alcocer 3B Ritteneria manca 344 4Z
Buiiol Ligerimys ellipticus 336 49
In the lacustrine and some pond-related localities, the eomyids have probably
been redeposited. In contrast to the glirids, the eomyids have been accumulated
largely in swamp and fluviatile deposits. A riparian, not aquatic habitat, closely
related to a high vegetation content may be deduced for the eomyids based on these
results. The interpretation of van de Weerd et al. (1978) and Daa:ms et al. (1984) of
the eomyids as indicators of wet environmental conditions is also strongly supported
by this result.
Additionally, we have studied two other variables, the molar ratio and the
metric variability, as indications of taphonomic differences among mMF As.
Other authors (e.g., Tobias, 1986) have paid some attention to the upper/lower
molar ratio of the fossil assemblages of vertebrates, which would indicate the rede-
posited condition of the samples having a low ratio. We have calculated this ratio for
the best represented species of our assemblages (table 4). The nearly constant ratio
around the 50% level shown by all the cases does not agree with the mentioned
hypothesis, since the preservation of several of these samples would indicate redeposi-
tion. Moreover, the conclusions of the study of Voorhies (1969) indicates that jaws and
skulls are both equally selected by currents. Our conclusion is that the frequent bias
in large mammals to higher numbers of jaws must be an artifact of sampling; jaws
have a better record because they are usually preserved with bone, but skulls are often
broken, and the isolated upper molars are more difficult to see.
471
Table 5. Variation Coefficient (CV: 100 X standard dev./mean) of the length of M1-2
of various Eomyidae and Gliridae species.
Locality Species N Var. cv
Gliridae
Moncalvillo Pseudodryomys ibericus 26 LM2/ 7.2
Santa Cilia Pseudodryomys ibericus 20 LM1/ 6.8
Moheda Pseudodryomys lopezae 28 LM1/ 4.9
Pseudodryomys lopezae 47 LM2/ 5.0
Pseudodryomys lopezae 25 LM/1 4.9
Pseudodryomys lopezae 39 LM/2 5.3
Eomyidae
Bergasa Rhodanomys transiens 245 LM/1-2 6.9
Moncalvillo Rhodanomys schlosseri 95 LM/1-2 6.8
Autol Rhodanomys schlosseri 38 LM/1-2 6.8
Santa Cilia Rhodanomys oscensis 195 LM/1-2 6.9
Quell Ritteneria molinae 30 LM/1-2 5.7
Cetina de Arag6n Ritteneria molinae 70 LM/1-2 6.0
Alcocer 3B Ritteneria manca 99 LM/1-2 6.2
Finally, we have studied the variability of some biometrical measurements, using

the Variation Coefficient (CV: 100 X st. dev./mean). Vianey-Liaud et al. (1986) have
used it as the main criteria to conclude the general taphonomic homogeneity of the
MFA. The CV of their samples varies between 1.83 and 12.2 for sample sizes from 14
to 220. Most authors have retained a CV between 4 and 10 for the most diagnostic
feature of species to consider probable taxonomic homogeneity of a sample (Simpson
et al., 1960; Sokal and Rolf, 1979), but there are no criteria of taphonomic homogene-
ity based on the CV. The procedure should be first to confirm it for many sites, then
to calculate the biometric parameters of the supposed homogeneous samples.
The best hypothetical estimate for the time of formation of basinal MFA ranges
from 25 years to 10,000 years (Behrensmeyer, 1978). Reworked specimens mixed with
younger assemblages must be separated by a longer interval (Koster, 1987), and conse-
quently the samples with accumulated-re deposited specimens mixed together with
reworked ones must vary significantly in some of its characters. Our CV shown in
table 5 does not show significant changes between localities. But it cannot demon-
strate that they have been formed during a similar time interval. The period of bone
accumulation and redeposition may be rather different without changes in the CV.
CONCLUSIONS
The preliminary results on the taphonomic factors affecting the mMFA forma-
tion have been based mainly on the study of the preservation state of micromammal
molars. After the procedure of Lacomba (1988), the preservation of glirids seems to
be in agreement with the sedimentologic al characteristics of the localities. Lacus-
trine and pond-related sites have the best preserved glirid teeth, swampy sites are
intermediate, and fluviatile sites have rounded, eroded specimens.
On the contrary, the preservation states of eomyids show an opposite distribu-

tion, the fluviatile and swampy sites having less eroded, transported material. These
472
differences are interpreted as a different way of production and burial in these
families. Eomyidae remains seem accumulated near water currents and are associated
with rich vegetation sites, as it was formerly assumed.
These differences in the taphonomic process emphasize that state of preserva-

tion does not depend on the sedimentary environment but depends on the taxa
involved. They also support the conclusion that predation has not been the principal
agent of production in these micromammal remains.
ACKNOWLEDGMENTS
This work is a part of the results of the research project numbers 2934-83 and
PB85-0022 sponsored by CICYT.
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Daams, R., 1976. Miocene rodents (Mammalia) from Cetina de Aragon (prov.
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Wetensch., B, v. 81(4), p. 448-473.
474
R.ELA'I10NS BETWEEN ~ALEOCUMATOLOGY AND
PUQ-PLEISTOCENE BIOSTRATIGRAPHIC DATA IN
WEST EUROPEAN COUNTRIES
Marie-Franfoise Bonifay
Laboratoire de Geologie du Quaternaire

C.N .R.S. - Luminy
Case 907
13Z88 Marseille Cedex 9, France
INTRODUCTION
The entire history of Plio-Pleistocene faunas is modified by the instability

typical of this period, which seems to explain most of the biostratigraphical difficul-
ties. In recent years, Plio-Pleistocene paleoclimatology has made tremendous
progress. Its scale has become shorter, especially for recent periods. Marine (isotopic
records) and continental data (sedimentology or palynology) have shown the existence
of short fluctuations of less than 10,000 years. These short climatic stages have very
little effect on the larger mammals and only influence the composition of associa-
tions. Medium length phenomena (100,000 years and more) correspond to transgres-
sion/regression marine cycles or to large-scale climato-sedimentary cycles such as
continental glaciations (WOrm or Valdai). We can see the influence of this scale on
migrations and species extinctions. These climatic phenomena interfere with evolu-
tion without really modifying it. In fact, we see only two major Plio-Pleistocene
events, which are Villafranchian fauna installation and Quaternary fauna migration
(Azzaroli, 1983).
The question is what is the true importance of major paleoclimatic events on the
global turnover of the larger mammals? Are correlations possible?
BIOSTRATIGRAPIDCAL DATA
The accepted succession of Plio-Pleistocene faunas in west European countries is

shown in figure 1. When Pliocene fauna disappear, Villafranchian fauna take their
place and are then in turn replaced by Quaternary fauna.
Villafranchian Fauna
The most important French sites are situated in the Massif Central (Bout, 1960)
(figure 3) whose history can be divided into two periods. The first (Inferior
Villafranchian fauna) shown by NM 16 (Etouaires) and NM 17 (Saint Vallier), corre-
sponds to increasingly rich associations. Many migrant species appear, such as
Leptobos, Equus, and Archidiskodon. The second (Superior Villafranchian fauna),
shown by the Seneze and Peyrolles sites, corresponds to poorer associations. It marks
the last appearance of the genera Nyctereutes, Anancus, Gazella, and of the Villa-
franchian~ Bovids, Panther, and Cervids species (Bonifay, 1983).

m.y. Fauna Biozones
16.1
z
']., Quaternary fauna
{evolved
r-
16.1
=
u
....
"'
archaic
....•..............•••••••..........•........
...a..
16.1 1 -
(Peyrolles)
VIllafranchian fauna (Sen~ze)
2 - inferior N.M 17 (St.Vallier)
16.1 N.M.l6 (Etouaires)

z 2,43
16.1
••••........•.•. ............•..............
...a..=
u ?
3
3,32
Pliocene fauna N.M.lS (Perpignan)
Fig. 1. Biostratigraphical succession in western European

countries.
Since 198Z, the age of the first French Villafranchian fauna has become clearer
through the work of the Clermont-Ferrand Laboratory (Guerppa, 1984). With the
exception of the Grand Combe whose fauna is poor and badly known, the following
groups have been identified (figure 4).
Perrier-Etouaires (Inferior Villafranchian fauna) is included in reworked fibrous

pumice sedimentation. Its age is between Z.47 and Z.35 Ma.
Perrier-Rocca Neyra, also Inferior Villafranchian fauna, but much later, is included
in the first inferior volcanic mudflow. Its age is between Z.35 and z.o Ma.
Pardines, the last Inferior Villafranchian fauna, is included in the first superior
volcanic mudflow. Its age is about Z.O Ma.
These three faunas were effected by Mont Dorian eruptions, which have been
situated between z. 7 and 1.6 Ma.
For Superior Villafranchian faunas, the Peyrolles site is included in the Allier
alluvial terrace which is situated between the beginning of the Sancy eruption
(1.Z Ma) and the Coudes trachyandesitic pumices whose age is 1.06 Ma.
From this new point of view, the following sites can be eliminated: Vialette in
France (Bandet et al., 1978), which is poorly known and whose fauna is dominated by
Tapirus, and Triversa in Italy (Arias et al., 1980). For the Triversa unit, whose age is
about 3.Z or. 3.1 Ma, Italian paleontologists think it is closer to the late Ruscinian
fauna from Peripignan than to the Montopoli Villafranchian fauna. For my part,
Triversa fauna cannot truly be considered to be a Villafranchian fauna.
Consequently, it can be thought that the first major faunal turnover, the instal-
lation of a French Villafranchian fauna, is later than was previously supposed and can
be situated at Z.4 Ma, after the Gauss/Matuyama event. ·
476
SPAIN FRANCE ITALY BIOLOGIC
EVENTS
AgeM.y. Massif Central and Mediterranean

peripheric areas areas
0,9 Soleihac Vallonnet Imola Leptobos and Villafranchian

Cervids disappear
1,0 Peyrolles Durfort
1,1 Ceyssaguet few fauna species
1,2 Villafranchian Lynx, Bovids

Panlher~
1,3 Sinzelles Farnetta rJrSt occurence of Hippopotamus
1,4 Tasso Canis event

last Mastodon and Gazelle
1,5 Seneze disappear
1,6 OlivoIa many Bovids
1,7 Nyctereutes disappear
11> --·--·-·-·····-··-··------------------------------------------------- -·-··-·········--··-··-·····-··

1,9 Le Coupet
2,0 La Puebla Pardines many Cer\'ids

de Valverde
2,1 St.Vallier
2,2
Perrier Rocameyra
2,3
2,4 Perrier Etouaires

1----------.Archidiskodotl
Equus
}
lirst occurcncc
2,5 Montopoli Leptobos
rtrSt Ml&odon ~fl.~~
2,6
Rincon I
2,7 - - - - - - - - -· - - - -· - - -
2,8
2,9
3,0 Tri\'crsa
3,1 Vialette
Fig. z. Succession in French, and neighboring countries, Villafranchian

fauna.
477
~---
j ClermonE-Ferrand
*Limoges _Q. "PEYROLLES;:::::: Lyon·
Mt.Dore~ :PARDINES_::::::-
sancy _.-- •PERRIER -
*
··~
........................
SENEZE "CEYSS~GUET
0VIALETTE
LE COUPET-.......•-....;. SINZEELES
~·!._UY
MEDITERRANEAN SEA
0 so IOOkm
L-----'---'
Fig. 3. Geographic situation of French Villafranchian sites.
478
Nature rl depolltll 111111 fallllll Abllolute age
0
1
2
3
4
s
"I Superior basalt from Chelx 0,58 ± 0,04 (R.671S)
6
7
i
8
0,83 ± 0,04 (FK.6926)
I
Pumices ofNeschen
9 F6 hippopotamus 11ernce
- Inferior basalt from Chelx 0,92 ± o,os (R.6757)
1,0
Trachyandesltlc pumices from Coudes 1,06 ±0,30 (8.91197)
1,1 FS Peyrolles ·
- Trachytlc pumices from Peyrolles
1,2 beginolng rl Sancy eruptloDS 1,2
1,3
1,4
1,5
1,6
1,7
1,8 t no deposit
1,9 J
2,0 ! . Superior tephra # 2,0
2,1 1 [!l.oubim"e de Pardlnes first superior volcsnic mudflow
~~.s
F3 Perrier Roccaneyra first inferior volcsnic mudflow
2,35 ± 0,06 (F.K.9409)
F2 Perrier~lnsubaeraleruptloo
2,4
2,5 Rh,mltlc fibrous uumk:es 2,47 ± 0,13 (F.K.8117)
"Grande Nappe" # 2,6-2,5
2,6
2,7
2,8
1,9
3,0 ruardelle grand combe

3,18 ± 0,09 (F.K.8799)
Fig. 4. Perrier plateau volcanic chronostratigraphy, based on Ly

Meng Hour, J.M. Cantagrel, A. de Goer de Herve, and P.M.
Vincent, 198Z; P. Mossand, J.M. Cantagrel, and P.M.
Vincent, 198Z; J .P .Poidevin,J.M, Cantagrel, and Guerppa,
1985. ( ) references to samples, L.A. 10 CNRS, Clermont-
Ferrand.
479
Quaternary Fauoa
After the late Villafranchian fauna such as those in Ceyssaguet or Peyrolles

(Haute-Loire) (Bout and Azzaroli, 1953) where a few species of Artiodactyls and very
abundant Equus stenonis (Perissodactyls) are present the Quaternary faunas appear
(Thouveny and Bonifay, 1984). In western European countries these temperate faunas
were subjected to climatic shifts linked to major continental glaciations known as
"Wiirm" in western Europe, as "Valda'i" in eastern Europe, and as "Wisconsin" in North
American countries. Western Europe acted as a ki.J;ld of container for temperate
species, while eastern Europe acted as a container for cold climate fauna, depending
on the dominant climatic component - whether cold or temperate - species expanded
from the northeast or from the southwest.
In our countries, we can divide temperate faunas into two groups. The first
group - temperate Quaternary faunas of the archaic type or transitional fauna
(Bonifay, 1978) is characterized only by Praemegacerids among the Cervids and is
accompanied only by the small Canis etruscus, between 1 and 0.5 Ma. The second
group - temperate Quaternary faunas of the evolved type is very close to modern
species and is characterized by Cervus elaphus accompanied by Canis lupus.
PALEOCLIMATIC DATA
Isotopic records are the most recent, the most continuous, and provide the best
results. Clearly, during the oldest period (figure 5) we can see a very distinct break at
about 3.2 Ma. The warm climate became a temperate climate, but the different
authors do not mention glaciation.
The first Pliocene glaciations took place before and after the Olduvai event
during the Matuyama reverse Chron (figure 6), with the most recent occurring during
the Brunhes Chron. The longest and the coldest glaciation is situated about 0.6 Ma.
COMPARISONS
Short Climatic Fluctuatious
Climatic isotopic records show that short fluctuations of less than 10,000 years
exist and that the same fluctuations occur in continental areas (figure 7). Using this
theory, Delpech (1983) drew a parallel between continental climatic stages (Laville et
Unstable temperate climate Warm climate
+3,0
...E
j~
+4,0
1
..,
Q)
..."'
Q)
.
a.
0 eQ)
""....
MI\TUYAMA GAUSS GILBERT
1,8 2,4 2,9 3,1 3,3
Fig. 5. Isotopic records, the oldest period, based on Shackleton .and Opdyke, 1977.
480
,.
0 ~
I...I
-{- -- - I? n
w
l
6 .c
.1 _j] l
Ill _______1. \
a
.....0
----.-1.0.. "'
~ n q
~
ltl
.....
"' ~
~-
;!i.e: z. __j]
0
ltl
.....
__u_ "' ~ n 1\
01
14 <
~
Ill
~ GG
_1 ~ J..i
16 ['..,
n 17
18 1'3 I0
10 <,
n 11
21 r:c:
~
l ~ "
"'
1-- ~')
.l_ n
n
_<;;_
- 0
- n
-
(
~
~
I? :
s
0
r
0
c:
<
)·
{.
)
~
/
~
- _...., ~ -
Fig. 6. Various climatic stages,
based on Shackleton and
Opdyke, 1976.
481
D© l.l':tlDtB~ ll'l\E
~
, ,c:
C)
f! ~
~ 0"' :ll
"',<::
)> (')0
il :!1
"' , c: (')
<: (')
0 ~<:
f?"'
C) )>
~~
"'
)>
';;
&' :ll
Sl
"'~
~
"' ..
5i "' ~"'
"" ..
~
"..
~
"' 3' ~~- 0
&l
~
~
R~
.. s· .. "'
~· ~
PHASES CLIMATIQUES 2' i;l "''!. ~ 0
(d'apres H . Laville)
~ 1\'
~ "
~-
~~
"'
!;
0
~ ~
II
--c;;.~.l
t sa'iga
~-- Sangl
,----
'INTERSTAOE' I II ·IV
XIV
XIII
1
RENNE
XII
XI I
X
IX
VIII
VII
VI
CERF
V Chevnuil
IV
Il l BCEUF· Sangliet
II RENNE
RENNE · BISON
Ct<EVAl
Fig. 7. Distribution of various species over the last 35,000 years in relation to
continental climatic stages of Laville, based on Delpech, 1983.
al., 1983) and fauna from Dordogne prehistoric sites over the last 35,000 years. These
climatic stages, either continental or isotopic in origin, were paralleled by Laville et
al. (1986). The authors gave further details, after having studied correlations between
the Grande Pile and the Les Echets pollen sequences.
For example, during isotopic stage Z and between 35,000 and 15,000 years, in the
continental western area, they distinguished an unstable short period consisting of I to
vn sediiJ1entologic stages, a threshold (stage Vlli) and then, between IX and XIV
stages, the installation of a cold climate with Rangifer tarandus faunas.
It is easy to verify whether the larger mammals can be perfectly correlated with
these short climatic fluctuations. In fact, they have very little influence on larger
fauna and change only the composition of association. Temperate species never
entirely disappear from our countries and they reappear immediately upon the return
of temperate conditions.
482
Medium-Size Climatic FluctuatioDS
Transgression/regression cycles in marine areas which correspond to major

climato-sedimentary cycles, such as glaciations in Alpine chronology (Wurm or Valdai),
give us medium-sized phenomena, which last about 100,000 years (Bonifay, 1980).
In western European countries, we can see that during the Brunhes Chron, and
despite several glaciations, in general the same temperate species reappear with the
exception of some subspecies. Only a few species become extinct and their extinction
does not correspond to a major paleobiologic event.
Major Faunal Turnover
In fact, in Plio-Pleistocene European faunas, there are only two major faunal
turnovers: the first, with the installation of the Villafranchian fauna at about Z.S Ma;
and the second, with the installation of Quaternary fauna at about 1.0 Ma.
However, we have seen that the climatic isotope curve shows two very unstable
periods after the 3.Z Ma crisis: the first near the Olduvai event; the second during the
Brunhes Chron.
I
l_ _,.._._ _ _l_ _ _ -=-
0 Adual raw. Return of temperate fauna
0,5
wd ~- od
unstable period
las~aciations
GG
cf ________________ _
secon turn over
relative stable period
DISAPPEARANCE
1,5 J superior
1___::':::_-----------
2 I
:>=
inferior
dominant temperate species
umtable period
first glaciations
temperate species installation
2,5 ------------ -- 'ffisi tuin-over-------------- stable period
DISAPPEARANCE
I
Qimatic crisis
warm fauna (warm-temperate)
3,5
Age biostratigraphic c6matic features isotopic record paleomag.
record of larger fauna
Fig. 8. Major continental turnover and major climatic crisis in isotopic
records: suggested correlations.
483
The first faunal turnover does not correspond to the climatic crisis 3.Z Ma ago
but could perhaps explain the disappearance of Pliocene faunas. In the continental
area, the first paleobiologic crisis about Z.4 Ma corresponds to the first Austrian
loesses at about Z.45 Ma, to the first Chinese loesses, and to the first cold cycle
between Z.4 and Z.3 Ma, but not to an unstable period of the isotopic curve which is
situated later (figure 8). These first glacial phenomena observed in the isotopic curve
near Olduvai correspond to the first disappearance of species from the Superior
Villafranchian faunas. The latest Villafranchian fauna and Quaternary fauna installa-
tion, which is the second major turnover, also took place during another stable isotopic
period which preceded the agitated Brunhes Chron.
CONCLUSIONS
The better climatic data now available - isotopic curves - does not explain
major biostratigraphic events. We can suppose that a correlation exists between other
fluctuations, such as volcanic events, but we should be able to identify them; or to
very dry periods such as those present in palynological studies, but would they be
sufficient, or to some paleomagnetic event.
However, we have found no correlation between major continental faunal turn-

over and major marine climatic events. Though it is difficult for a scientific work to
end on a problem, for the present study a simple question remains as to the lack of
concordance observed between the different phenomena and that question is: Why?
REFERENCES
Arias, C., Azzaroli, A., Bigazzi, G., and Bonadonna, F.P., 1980. Magnetostratigraphy
and Pliocene-Pleistocene boundary in Italy. Quaternary Research, v. 13, p.
65-74.
Azzaroli, A., 1983. Quaternary mammals and the end Villafranchian dispersal event -
A turning point in the history of Eurasia. Palaeogeography, Palaeoclimatology,
Palaeoecology, v. 44, p. 117-139.
Bandet, Y ., Don ville, B., and Michaux, J ., 197 8. Etude geologique et g~ochronologique
du site villafranchien de Vialette (Haute Loire). Bull. Soc. Geol. de France, v.
ZO, P• Z45-Z51.
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Guerppa, 1984. P:resence possible d'Hominides en Auvergne au Pliocene superieur (Z,5
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484
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de l'Ass. franC~• pour l'Et. du Quatern, v. 1/Z, p. 35-46.
Ly Meng Hour, Cantagrel, J.M., de Goer de Hervli, A., and Vincent, P.M., 198Z.
Revision tephochronologique des depots fossiliferes plio-pleistodmes des
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Shackleton, N.J. and Opdyke, N.D., 1976. Oxygen isotope and palaeomagnetic stratig-
raphy of Pacific core V. Z8-Z39. Late Pliocene to latest Pleistocene. Geological
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evidence from early northern hemisphere glaciation. Nature, v. Z70, p. Z16-Z19.
Thouveny, N. and Bonifay, E., 1984. New chronological data on European Plio-
Pleistocene faunas and hominid occupation sites. Nature, v. 308, p. 355-358.
485
SMALL MAMMAL TAPHONOMY
Peter Andrews
British Museum (Natural History)

London, England
INTRODUCTION
The study of small mammal faunas presents particular problems for the verte-
brate paleontologist. This is because small mammals are strongly affected by
taphonomic processes that alter the original composition of the faunal community.
Particular species may be selected for or against, with the result that while some
species may be missing altogether from a fossil assemblage, even though they were
present in the community from which the fossil assemblage was derived, other species
may be present in numbers unrelated to their original abundance. This makes it diffi-
cult to place too much reliance either on species presence or absence or even more on
the relative abundances of species which might be used as the basis for paleoecolog-
ical reconstruction.
The processes operating during the preservation and fossilization of small

mammals are shown in figure 1 (Andrews, 1989). The living community forms a death
DEATH
DEATH)
MODIFIED SHORTLY AFTER DEATH

(BY HUMANS/SCAVENGERS/TRAMPLING)
MODIFIED BEFORE/DURING BURIAL

(WEATHERING/TRANSPORT/SYNDIAGENESIS)
FOSSIL ASSEMBLAGE
Fig. 1. Stages in the formation and modification of small mammal bone assem-
blages, from living animals to fossilize~! bones. Formation stages are shown
on the left, and modifications that can occur from one stage to the next are
shown on the right.

Edited by E.H. lindsay et al.
assemblage of animals derived from it, either without alteration or biased by the
cause of death, in the case of small mammals usually predation. The death assem-
blage in turn gives rise to a bone accumulation, usually further altered by processes of
decay and scavenging activity. The bones left exposed are then subjected to a variety
of processes both above ground, such as trampling by the still living members of the
animal community, weathering by sun, rain, and wind, and transport by wind or water;
and below ground, by soil compaction, soil corrosion, root etching, and sometimes by
secondary transport of soil or sediment. Some, none, or all of these processes may act
sequentially to alter the original composition of small mammal communities, and an
understanding of the extent of modification is necessary before reaching conclusions
as to the presence/absence of particular taxa, or interpreting the paleoecology of the
fauna. Some of these factors will be briefly reviewed here based on detailed results
being presented elsewhere (Andrews, 1989).
PREDATION
Predation is probably the single most significant factor in the accumulation of

small mammal remains. It is a significant cause of death in most small mammal
communities, but, even more, it is probably the most important single agent respon-
sible for producing concentrations of small mammal remains. Remains of pellets from
nocturnal or diurnal birds of prey yield enormous quantities of bone (figure Z) and in
certain cases scat concentrations from mammalian carnivores do likewise. Small
mammal prey accumulations, however, are all biased according to the hunting style
and abilities of the predator concerned. This may be by size of prey, for all predators
hunt prey within certain size limits, or by species preference; for example, microtines
are favored prey for many predators, or by activity patterns of the prey, whether
nocturnal or diurnal, and many other factors. 1
Fig. z. Example of the bones from a broken down

pellet accumulation below a Barn owl's nest.
The bones are mixed with the stony soil, and
already the degree of breakage is greater
than would have been present in the intact
pellets.
488
50
40
UK average
MNI-47,865
30 N- 10/14
20
~ 10
..!!
Q.
- 0
. .,.. •. •..••
! ..: 'ii!
>-
! ~
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:
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~
';:.
30
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.::
Q
II) Stratton, Dorset
40 Q
MNI-147
00:
N -8/14
50 . -.
-·•....e!
:!li
Fig. 3. Barn owl diet in Britain. The histogram shows

percentage prey representation in the diet,
with the U.K. average shown above the line
and a single pellet accumulation from one
area of southern England shown below the
line. The minimum numbers of individuals
(MNI) are based on skull and mandible
counts. The numbers of species present (N)
are shown compared with the theoretically
possible number of small mammal species (14)
available for consumption by the Barn owl.
The effects of this predator selection can be illustrated with a simple example.
In figure 3, the prey of the Barn owl is shown, comparing the U.K. average prey for
this species above the line, based on nearly 48,000 prey records (Glue, 1967), with a
single pellet accumulation below the line with only 147 prey individuals. In the U.K.
as a whole, the Barn owl takes 10 species out of a possible 14 (excluding the House
mouse), rarely or never taking such animals as rats, squirrels, or hedgehogs. Even on
this broad spectrum, therefore, the Barn owl only takes 71% of the possible small
mammals available to it. When a single pellet sample is considered, sampling error
reduces the prey spectrum still further, since species eaten only rarely will be less
likely to be represented than common prey species. In the example here, only eight
species are present in the owl assemblage, 80% of the country-wide sample and now
only just over 50% of the possible prey.
The Barn owl was chosen for this example because it is an intermediate feeder,
neither very specialized nor very opportunistic. Highly selective predators, such as
the vole specialists (e.g. Strix nebulosa), produce prey assemblages even more limited,
while opportunistic predators such as the owls Bubo bubo and Strix aluco or the diurnal
raptor Buteo buteo produce prey assemblages representing up to 85% of the small
mammal communities available to them. For all predators, however, there is this
489
Fig. 4. Digestion of small mammals by predators:
A. digestion of the tip of a microtine incisor
by a Long-eared owl (Asio otus); B. digestion
of a soricid molar by a Hen harrier (Circus
cyaneus); C. digestion of a microtine distal
humerus by a Red fox (Vulpes vulpes).
combination of selectivity and sampling error which ensure that species present in the
original community are not represented in the fossil record, and some knowledge of
the type or species of predator will provide an indication of the potential size and
nature of the predator bias.
New approaches to small mammal taphonomy are atttempting to provide a

framework against which the effects of predator bias can be estimated (Mellett, 1974;
Mayhew, 1977; Korth, 1979; Dodson and Wexlar, 1979; Andrews and Evans, 1983;
Andrews, 1989; Kowalski, in press; Fernandez-Jalv o, in prep.). This entails the evalua-
tion of predator and other modifications on recent bone assemblages, and using the
ensuing pattern of modifications to recognize predator activity on assemblages of
unknown origin. This is a straightforward process for recent bone assemblages, and
predator activity and type can be recognized with a high degree of success (Andrews,
1989), but with older fossil assemblages there are some additional problems to over-
come. One of these is that the predator itself may be an extinct species producing a
prey assemblage not comparable to that of any extant predator. Another even more
difficult problem is that secondary modification resulting from later stages of
taphonomic process (figure 1), especially the results of burial and fossilization pro-
490
buzzard
I ' ' ' ' I Ill ' fl Ill I II I I I' 111f111 I f1111f1111JIIIIJIII 'I'"' IIIII
10 20 30 40 50 MM
II j I
1 [., ; ,
I I I 'I
• 1 I ~ 1 I , •
I I
I I
Fig. 5. Breakage of lagomorph mandibles and teeth by (above) a

Common buzzard (Buteo buteo) in Wales and (below) a
Coyote (Canis latrans) in North America.
cesses, may so alter the bone assemblage that the original predator-induced pattern
can no longer be recognized.
The most diagnostic category of predator-induced modification of small mammal

bones is seen in the effects of digestion. This produces enamel loss from the teeth,
with extreme digestion also affecting the dentine, and the patterns of modification
differ from one prey species to another. Digestion of rodent incisors varies less than
that of molars, with their more greatly differing morphologies. And, digestion of
rodent teeth generally differs from that of insectivores and lagomorpha. When the
evidence of digestion from all taxonomic groups of prey. is combined, consistent
patterns emerge which are specific to individual predator species and which serve to
identify the predator. Some examples of digested small mammal bones and teeth are
shown in figure 4.
Breakage of small mammal bone is also diagnostic in undisturbed pellet or scat

assemblages, although there are problems when secondary breakage occurs (see
below). Owls and diurnal raptors eating small prey individuals produce little breakage
of the bones of their prey, but a much higher degree of breakage occurs when prey
size is large relative to the size of the predator. Mammalian carnivores habitually
break up the bones of their prey, producing distinctive patterns of bone element
491
preservation and breakage (Andrews and Evans, 1983). Some examples are shown in
figure 5.
POST-PREDATOR MODIFICATIONS
A number of processes may operate on small mammal accumulations to further

alter them (figure 1). Some are less important than others, not so much because of
their frequency of occurrence but because they are so destructive as to completely
destroy the assemblages. Water transport, for instance, readily disperses small
mammal bone, which may become rounded and broken in the process, but it is not
likely to produce great concentrations of bone. Small pockets of bone may occur
through this agency, but nothing comparable to predator accumulations. The rounding
produced by attrition is similar to that produced by some mammalian carnivores
during digestion of small mammal bone, but attrition will either produce rounding
without digestive corrosion, if none had previously been produced by a predator, or, if
corrosion is already present, the rounding would be obviously superimposed on it and so
distinguishable. A useful association between kinds and degrees of modification with
sedimentary environment has been made by Alvarez-Sierra (Chapter Z6 in this
volume), by which she showed an apparent association between energy of water flow,
as indicated by particle size and type of sediment, and the degree of rounding of the
bones.
Weathering is also a destructive process of small mammal bone. Current obser-

vations show that weathering produces splitting and cracking of small mammal teeth
and breakdown of thin bone on skulls and postcrania after two years exposure (figure
6), and weaker elements such as scapula and ribs are quickly destroyed. Soil corrosion
after burial produces a similar effect, although distinguishable from weathering by the
extensive pitting that occurs of both teeth and postcrania. Corrosion occurs in a
variety of soil types, strongest in acid soils but present also in alkaline soils on chalk
and limestone, where it is apparently the organic acids present in organic-rich soils
that produces it. Soil corrosion can be distinguished from the effects of digestion by
its more extensive but patchy distribution over the bones and teeth (figure 6), and
weathering can be distinguished by the splitting of both bones and teeth. Although
their manner of modification of the surfaces of the bones is different and distinguish-
able from the effects of digestion, if either weathering or soil corrosion is super-
imposed onto digested bone, it then becomes difficult to identify the source of the
digestion, since it is the already altered parts of the bone that are affected most
strongly by the later processes. Some examples of the modifications are shown in
figures 5 and 6, which are extracted from the much fuller account provided by
Andrews (1989).
The process of destruction of small mammal bone is continued during fossiliza-

tion. Sediment movement and compression usually ensures the removal and destruc-
tion of all but the strongest elements, but there are no records available to show how
different species are affected by this process. Isolated teeth are the most commonly
preserved element in most fossil assemblages because of their small size and durabil-
ity, and it would seem likely that even small differences in size and shape could
produce big differences in survivability of teeth from different taxa.
CONCLUSIONS
This brief survey of current work into small mammal taphonomy can only point
to problems rather than attempt to provide answers. Our work on small mammal
faunas in caves has succeeded in identifying several of the predator species respon-
sible for accumulating the rich bone accumulations present at different parts of the
Pleistocene sequence of Westbury Cave (Andrews, 1989). This has been particularly
useful, for instance, in identifying an apparently major faunal change in successive
levels as being due to taphonomic rather than paleoecologic causes. At another level
of the same cave, weathering has been found to be responsbile for the in situ breakage
492
Fig. 6. Weathering damage on the occlusal surface of a
microtine molar (above) and soil corrosion of a tibia
shaft (below).
of much of the bone. Specific predator action has also been identified in the Pleisto-
cene succession of another cave in Spain and, at some levels, this is combined with soil
corrosion of a particular type found in limestone caves (Fernandez-Jalvo, in prep.).
Predator action of an unspecified kind has also been identified on the small mammal
bones from Miocene deposits of Pa~alar, Turkey (Alpagut and Andrews, Chapter 25 in
this volume), but here the later transport and breakage of bone has been so great as to
limit predator identification to a general categorization of predator type, although
this is sufficient in itself to indicate the probable loss of very small species (less than
1 00 g) from the record.
There is clearly great potential benefit in the identification of taphonomic bias

in small mammal faunas. We have made only the most preliminary beginnings to this
study, which should be a part of every faunal analysis undertaken. In most of the
papers published in this volume, little attention has been paid to taphonomic effects
on the fauna, even when great weight is being put on the first appearances or dis-
appearances of species in the fossil record, or on the significance of faunal associa-
tions. The question must always be asked: whether the evidence is a true reflection
of the past or whether it has been altered in some way by taphonomic modifications.
Only when this has been done can the evidence be accepted as valid.
REFERENCES
Andrews, P., 1989. Owls, Caves and Fossils: Predation and Preservation of Small
Mammals in the Fossil Record. British Museum (Natural History), London.
493
Andrews, P. and Evans, E.M.N., 1983. Small mammal bone accumulations produced by
mammalian carnivores. Paleobiology, v. 9, p. 289-307.
Dodson, P. and Wexlar, D., 1979. Taphonomic investigation of owl pellets. Paleo-
biology, v. 5, p. 275-284.
Glue, D., 1967. Prey taken by the Barn owl in Britain and Ireland. Bird Study, v. 14,
p. 169-183.
Korth, W.W., 1979. Taphonomy of microvertebrate fossil assemblages. Ann. Carnegie
Mus., v. 48, p. 235-285.
Kowalski, K., in press. Some problems of the taphonomy of small mammals.
Mayhew, D.F., 1977. Avian predators as accumulators of fossil mamYlal material.
Boreas, v. 6, p. 25-31. ·
Mellett, J.S., 1974. Scatological origin of microvertebrate fossil accumulations.
Science, v. 185, p. 349-350.
494
HIPPARION DATUM AND ITS CHRONOLOGie EVIDENCE
IN THE MEDITERRANEAN AREA
Sevket Sen
Laboratoire de P&~leontologie des Vert~bres

Universite Pierre et Marie Curie
U.A. 720 DU CNRS
4 place Jussieu
75Z5Z Paris Cedex 05, France
INTRODUCTION
For mammalian biochronology in Europe, the immigation of hipparionine equids

into the Mediterranean and central European areas constitutes a major event, which
characterizes the Aragonian-Vallesian boundary (Fahlbusch, 1976; Ginsburg, 1975).
Berggren and van Couvering (1974) popularized this event by introducing the concept
of "Hipparion Datum."
During the late 60s and the 70s, the generally adopted opm10n was that this
group had a high rate of expansion, and that it invaded the whole of Eurasia and North
Africa almost instantaneously with respect to resolution of geologic time. Berggren
and van Couvering (1978, p. 49) restated this opinion as: "The real time difference in
the observed first appearance datum in this vast region is probably less than 0.5
m.y." However, hypotheses on the synchrony of Hipparion immigration throughout
Eurasia and the Mediterranean area were founded on few chronological data. One can
even wonder whether there exists anywhere suitable time data for such an hypoth-
esis. The idea of synchrony is partly due to the limits of biochronological resolution of
large mammal faunas. In fact, Hipparion is the unique American immigrant among the
Eurasian large mammals of the early late Miocene.
Ages ranging between 1Z.5 and 10 Ma were proposed for the first occurrence of
this group in Eurasia and the Mediterranean area. Berggren and his collaborators
(Berggren and van Couvering, 1974, 1978; van Couvering and Berggren, 1977; Berggren
et al., 1985) argued for an age of 1Z.5 Ma or between 1Z.5 and 1Z Ma for this event,
based on some radiometric data from Howenegg and Bou Hanifia. However, radio-
metric datings and magnetostratigraphic correlations on the continental deposits of
Turkey and the Indian subcontinent provided some younger ages for the oldest
Hipparion faunas of these countries. It appears, therefore, that the synchrony of this
event as Berggren and his collaborators understand can no longer be accepted in such a
large area. Their assumption does not take into account ecological and paleogeo-
graphical factors which influenced equid dispersal. The aim of this paper is not to
insist more on this subject, but to present a review of available data on the Hipparion
Datum, with some new results (K/ Ar dating and magnetostratigraphy) recently
obtained from some Hipparion localities in the Mediterranean area.

Fig. 1. Some Vallesian mammal localities with Hipparion in the
Mediterranean area. 1. Howenegg, z. Hostalets de Pierola, 3. Bou
Hanifia, 4. Kastellios Hill, 5. Esme-Akcakoy, 6. Bayraktepe n,
7. Yassioren. Some of these localities also contain small
mammals.
HlPPARION IMMIGRATION
The earliest occurrence of the first North American hipparionine equid

CormohipParion goorisi was calibrated as being 15 :J: 0.5 Ma old (Woodburne et al.,
1981; Flynn et al., 1984). It is at present obvious that this group emigrated into
Eurasia much later. In several studies on the New World and the Old World
hipparionines, a common interpretation is that the ancestor for the Eurasian species
lived in North America. The potential ancestor is, according to Woodburne et al.
(1981), Cormohipparion sphenodus (Cope, 1889), known in North American sequences
between 13.6 and 11 Ma. On the other hand, Bernor and Hussain (1985) argued that
Cormohipparion occidentale is particularly similar to the European species Hipparion
primigenium in its morphological features. In any case, as demonstrated by Bernor
and Hussain (1985) (see also Bernor et al., 1987a,b), the Old World hipparionines
probably have a monophyletic origin.
The generally accepted opinion was that this emigration happened some 1Z.5 Ma
ago. Woodburne et al. (1981) explained this event as a result of a major drop in sea
level to about 80 m below the present-day level, between 11 and 1 Z Ma, and this drop
gave rise to a land connection across Beringia. I have reproduced in figure 3 the most
recent curve of eustatic fluctuations (Haq et al., 1987; fig. Z) concerning the period
from 16 to 9 Ma. It appears that sea level drop was progressive from a high level at
about 15 Ma to a sharp fall a little before 10 Ma. This time interval also contains
three successive fluctuations, with a general tendency toward decrease in sea level.
The curve seems to speak in· favor of a land connection at about 11 to 10 Ma between
North America and Asia. For a reasonable correlation of the sea level drop with
Hipparion emigration from North America to Eurasia, it would be better to have some
calibrated key-localities in Asia. Unfortunately, no valid absolute ages have been
published until now on the oldest Hipparion of Asian localities, except in the Indian
subcontinent where the Hipparion occurrence is apparently much later than in the
496
other parts of Eurasia. Detailed magnetostratigraphic studies in the Potwar Plateau
area (Pakistan) led to the calibration of the first occurrence of Hipparion in the
Siwalik sequences as 9.5 Ma (Barry et al., 1985). It should be noted that this age was
obtained by the correlation of magnetic polarity zones with the geomagnetic polarity
time scale (GMPTS) of Mankinen and Dalrymple (1979). The correlation of the same
polarity successions with more recent GMPTS will increase this age by some 0.3 Ma.
BIOCHRONOLOGIC BASIS OF HIPPARION DATUM
As initially defined at the 1975 Munich Symposium (Fablbusch, 1976), European

faunas with Anchitherium and without Hipparion belong to the Aragonian, and the
Vallesian begins with the occurrence of Hipparion. Concerning small mammal associa-
tions, the appearance of Progonomys is equivalent to that of Hipparion. However, it
should be noted that Progonomys appears, in the Vallesian type area, somewhat later
than Hipparion. Other rodent taxa have been more or less associated with this bound-
ary, such as the Cricetodon-Hispanomys lineage, Cricetulodon and Megacricetodon
(Agusti et al., 1984; Unay and de Bruijn, 1984).
Several localities in Europe and North Africa have yielded so-called early
Vallesian Hipparion (figure 1). The aim of this paper is not to discuss what generic
name should be employed for the Vallesian forms of the Old World nor which particu-
lar horse association presents the more archaic characters than the other. These
subjects were debated by several authors (Forsten, 1968; Sen et al., 1978; Steasche and
Sondaar, 1979; Bernor et al., 1980, 1987b; Woodburne et al., 1981) who brought some
light on the early Vallesian horses of the Old World. Woodburne and Bernor (1980)
defined some superspecific groups for the Old World hipparioniones; this paper only
concerns the Group 1 species. Bernor et al. (1987a, p. 44) noted that the Howenegg
Hipparion primigenium is "the most primitive species" of the Old World hipparionines
compared with H. catalaunicum from Spain and H. africanum from Algeria.
According to these studies, and taking into account chronologie data on the
earliest Old World species, some localities present more interest than others for a
biochronologic definition of the Aragonian-Vallesian boundary. These are Hostalets de
Pierola in Spain, Howenegg in Germany, Bou Hanifia in Algeria, Kastellios Hill in
Greece, Esme-Akcakoy, YassiOrE;'n, and Bayraktepe n in Turkey (figure 1).
In the type areas of Aragonian and Vallesian, this boundary is not characterized
by any particular event, e.g. faunal extinctions at the end of the Aragonian. Moy~
Sola and Agu~t( (1985, p. 403) note that in the Valles-Penedes Basin "this limit does
not represent an important faunistic change and is only marked by the arrival of
Hipparion ... from this point .of view the other limits are qualitatively more important,
especially the MN 9/MN 10 limit." Figure Z illustrates the time range of some char-
acteristic rodent taxa and shows that no particular event occurred when Hipparion
appeared in the Valles-Penedes Basin.
On the other hand, the Hipparion occurrence is apparently diachronous in differ-

ent basins of the Iberian Peninsula. Lopez Martinez et al. (1986) and Alvarez Sierra et
al. (1987) showed that Hipparion appears somewhat later in the Duero Basin than in
the basin of Valles-Penedes and Calatayud-Daroca. This diachrony is stated on the
correlation of rich rodent associations from these basins.
In another area, namely in Turkey, the faunal succession does not show any
significant change in relation to the Hipparion arrival. According to the data pub-
lished by Unay and de Bruijn (1984) on the Dardanelles localities and with my own
observations on some other Turkish localities, changes in the composition of mammal
faunas occurred either before the first occurrence of Hipparion (replacement of
Turkomys by Byzantinia, Megacricetodon extinction, appearance of Cricetulodon) or
following it (the first Progonomys).
497
5 p A l N T u R KE y
Valles-Penedes
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HIPPARION DATUM PLANE Yassioren
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.
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Fig. z. Time ranges of some rodent genera associated with the Hiwarion Datum in
the Valles-Pene!les Basin (Spain) and in Turkey.
Faunal composition for the Turkish localities mentioned in figure Z are given by
Unay (1981), Sickenberg et al. (1975), and Sen (1986). Yassioren and Esme-Akcakoy
faunas possess several common genera and species including Hipparion senyureki
Ozansoy, 1965. Common opinion among paleontologists who studied these faunas
(Steasche and Sondaar, 1979, described the hipparionines) is that they are the oldest
Vallesian localities of Turkey. The locality of Yassioren also yielded some remains of
small mammals: Schizogalerix sp., Proochotona sp., Byzantinia dardanellensis,
Cricetulodon hartenbergeri, Atlantoxerus sp., and Myomimus dehmi (Sen, 1986). This
assemblage is quite similar to that of Bayraktepe n, which also contains a still
undescribed Hipparion. Unay (1981) and Unay and de Bruijn (1984) attributed an age of
middle-late Vallesian to the Bayraktepe n fauna. If Progonomys is absent at
Yassioren, its presence at Bayraktepe n is proven by a single first upper molar.
In figure Z, the time ranges of some rodent genera from Turkish localities are
tentatively compared with their equivalents from the Valles-Penedes Basin. In both
areas, Cricetulodon appears in the faunas without Hipparion, while Progonomys
appears after the Hipparion arrival. However, the extinction of Megacricetodon is
earlier in Turkey than in Spain. In the same way, the genus Turkomys in Turkey is
replaced by Byzantinia far earlier than in Spain if we consider Cricetodon and
Hispanomys as their western equivalents.
Unfortunately, none of the mentioned Turkish localities, nor the localities in the
Valles-Pene!les Basin and in other Spanish basins, have associated radiometric or
paleomagnetic dates. Thus, there are no relevant absolute ages for the first Hipparion
occurrence in Spain and in Turkey.
498
RADIOMETRIC DATA lN EUROPE AND NORTH AFRICA
I have to emphasize first the rarity of radiometric ages relevant to the

Hipparion Datum. In spite of this situation, some scarce data help us to delimit the
age of this event and to understand why different ages were proposed for the
Hipparion Datum in Europe and North Africa. The age of 1Z.5 Ma (or between 1Z.5
and 1Z Ma) proposed by Berggren and van Couvering (1974, 1978), and later repeated
.by several authors, is radiometrically founded on data from two localities: Howenegg
in West Germany and Bou Hanifia in Algeria.
The first important locality, Howenegg in southwest Germany, yielded two

radiometric dates: 12.4 ± 1 Ma on hornblende of the fossiliferous level tuffs (Lippolt
et al., 1963) and l0.8 ± 0.4 Ma on whole rock analysis of a basaltic bomb found within
the fossiliferous bed (Baranyi et al., 1976). In a letter to J.D. Becker-Platen (Becker-
Platen et al., 1977, p. 164), H.J. Lippolt acknowledges that, concerning the first date,
"it is particularly difficult. to prove if this age is true, because the genesis of horn-
blende was not explained." H. Tobien (1982, 1986), an authority on Howenegg stratig-
raphy and its fauna, preferred the second date as the age of this fauna.
The Bou Hanifia continental deposits in northwest Algeria yielded further

determinations bearing on the Hipparion Datum. A simplified stratigraphy of these
deposits in the vicinity of Bou Hanifia Dam - where are situated the Bou Hanifia
mammal localities- is as follows:
1. Marine Anaseur Formation: Gray-green or blue marls near the top of this forma-
tion contain planktonic foraminifera attributed to the zone N 13 (Ouda and Ameur,
1978) and nannoplankton which indicates a correlation near the NN 6/NN 7 bound-
ary (Ameur et al., 1979).
z. Continental Bou Hanifia Formation: It overlies the Anaseur Formation with an

erosional unconformity and develops a thickness of 300 m or more around the dam
lake.
3. Unnamed upper marine deposits: The Bou Hanifia Formation is unconformably

overlain by marine deposits from which Ouda and Ameur (1978) mentioned a plank-
tonic foraminifera fauna of the late Tortonian.
Mammal localities as well as K/ Ar datings are known from the Bou Hanifia
Formation. It is an alternation of conglomeratic, sandy, and marly sediments of
grayish, pinkish, or reddish colors. In the lower part of this formation, Ouda and
Ameur (1978, p. 415) mentioned the presence of marine intercalations with planktonic
foraminifera that might indicate N 15 zone. During 1981 fieldwork, it was not possi-
ble to relocate these levels and to confirm this observation.
Sediments in the lower half of this formation contain three thin volcanic ash
levels. Chabbar-Ameur et al. (1976) published a K/ Ar date from one of these levels
without noting precisely its stratigraphic position within the formation. This date is
1 Z.18 ± 1.03 Ma. In 1981, I collected samples from the first volcanic ash level (thick-
ness about ZO em) observed 11 m above the base of this formation. New K/Ar dating
realized by P.Y. Gillot at the Centre des Faibles Radioactivites (Gif sur Yvette) on
sorted biotites yields an age of 1Z.03 ± O.Z5 Ma. This new date is in good agreement
with the previous one, and it may show that the volcanic ash in the sediments of the
Bou Hanifia Formation were issued from the same volcanic activity; its occurrence in
different levels may be interpreted as a product of redeposition.
These two K/ Ar dates indicate the oldest age that can be attributed to the Bou
Hanifia Formation. However, they cannot be extrapolated to represent the age of the
mammal-bearing horizons, which are situated near the top of the formation and some
100 m above the ash levels.
499
Several K/Ar dates from the middle and upper Miocene continental deposits in
Turkey led Becker-Platen et al. (1977) to propose an age between 10.8 and 11.1 Ma for
the Aragonian-Vallesian boundary, in other words for the Hipparion Datum. Among
the mammal localities dated by these authors, Yenieskihisar 1 (13.1 ± 0.5 Ma) yielded
a rich late Aragonian fauna, while the poor fauna of Yenieskihisar 2. (11.1 ± 0.2. Ma)
can belong to Aragonian as well as to Vallesian. At the locality of Yaylacilar, the
mammal-bearing horizon without Hipparion is situated 2.0 m above a tuff layer which
is dated at 11.6 ± 0.2.5 Ma. Localities with Hipparion yielded K/Ar dates younger than
10 Ma. Becker-Platen et al. (1977) proposed an age between 10.8 and 11.1 Ma for the
Hipparion Datum, taking into account the fact that this group is always found when
present in a fauna.
From all these K/ Ar dates, it can be concluded that no evidence exists on the
presence of hipparionine horses in Europe and North Africa older than 12. Ma.
MAGNETOSTRATIGRAPmC DATA
The best magnetostratigraphic calibration of the Hipparion Datum was done in

the Siwaliks of Pakistan. The first Hipparion occurs, in the Nagri Formation, in the
lower part of a long normal polarity zone which is accurately correlated with Chron 9
(Anomaly 5 equivalent). As mentioned above, Barry et al. (1985) attributed to this
event an age of 9.5 Ma.
In the Eastern Paratethys area (Moldavia, Ukraine, and Caucasus), the earliest
Hipparion occurs in the Middle Sarmatian. According to Chepalyga et al. (1985, P•
137) "the Gritsevian complex with the oldest Hipparion primigenium ... is character-
HIPPARION DATUM EUSTATIC CURVES
INDIAN
SUBCONTINENT
MEDITERRANEAN
AREA
Berggren &
Van Couverin~r 1974 ~
NORTH
AMERICA Woorihurne et al. 1981 ~
16
Fig. 3. Hipparion Datum in North America, Indian subcontinent, and

Mediterranean area, and its relationships with the sea level fluctua-
tions (after Haq et al., 1987). Chron numbers are from LaBrecq et
al. (1977). The Chron 9 of this figure is equivalent to Chron 11 in
the more recent terminology of Berggren et al. (1985).
500
VG P latitude Ma
I ) 8,49
8,80
8.98
-
~--
10.30
10.43
10.48
.,.
.,. " "
/
10.91
10.99
?
11.47
[m I
11,63
11.77
7 12.03
I
I
i I GMPTS
i
Lz
I Harland et al. till
I
-90 +90
Fig. 4. Magnetostratigraphi c study in the upper part of the Bou
Hanifia Formation. From left to the right, lithology of the
sampled section, stratigraphic position of the mammalian
localities BH 1 and BH 5, paleomagnetic sites, latitudes of
the virtual geomagnetic pole (VGP) in each site, succession
of the polarities in this section, and their correlation with
the geomagnetic polarity time scale of Harland et al. (198Z).
ized by the negative polarity of the upper 10 paleomagnetic epoch (10.8-10.3 Ma). So
the first hipparions appeared near 11 Ma." In a more detailed paper, Pevzner et al.
(1987) gave the results of their magnetostratigraphi c studies in Moldavia, and proposed
the correlation of the normal polarities obtained in the sections of Bujor 1, Kalfa,
Lapushna, and Atavaska with Chron 11. These localities yielded H. sarmaticum.
Chepalyga et al. (1985) consider these localities correlative with Chron 9. In fact, in
the sections studied by Pevzner et al. (1987), the polarity succession is poor (short
sections and with only one polarity zone), and these results cannot be correlated
securely with the GMPTS.
501
The Kastellios Hill section in central Crete yielded, from several levels, remains
of small and large mammals, and it also contains some marine intercalations with
planktonic foraminifera (de Bruijn et al., 1971; de Bruijn and Zachariasse, 1979).
Among the mammalian species, I have to mention Hipparion sp., compared by these
authors to H. primigenium, and two species of Progonomys. The mammal associations
of Kastellios Hill are attributed to the Vallesian, as being close to the MN 9/MN 10
zone boundary. The planktonic foraminifera indicate the base of N 16 zone (de Bruijn
and Zachariasse, 1979).
Detailed magnetostratigraphic study of an 88 m thick section in Kastellios Hill

(Sen et al., 1986) yielded reversed polarities which are interrupted by two short normal
zones, each 1 m thick. Taking into account the biochronologic data from several
levels (small and large mammals, planktonic foraminifera, ostracods, pollen), this
polarity succession was correlated with a part of Chron 10. This correlation confers
on these faunas an age situated between 10.3 and 11 Ma.
In the Bou Hanifia section in Algeria, I sampled 37 sites in the upper part of the
Bou Hanifia Formation (see above regarding the stratigraphy). The sampling is not
continuous and it is interrupted by two major gaps due to the poorness of the out-
crops. The locality BH 1, which has yielded among other mammals Hipparion
africanum and Progonomys cathalai (Jaeger et al., 1973), is situated in the lower gap.
The stratigraphic position of the small mammal locality BH 5 is reported on figure 4.
This figure also presents the stratigraphy of the sampled section, latitudes of the
virtual geomagnetic pole (VGP) in each site, the succession of the magnetic polarities,
and their correlation with the GMPTS of Harland et al. (198Z) (see details of the
paleomagnetic study in Sen, 1986). According to these results, the small mammal
locality BH 5, which also yielded Progonomys cathalai, should be as old as 10.3 Ma,
and the age of the Hipparion locality BH 1 should be a little older than 10.5 Ma, but
certainly younger than 1 Z Ma.
In East Africa, the magnetostratigraphy of the Ngorora Formation has been

studied at the type Kabarsero section (Tauxe et al., 1985). Bishop and Pickford (1975)
have mentioned the Hipparion occurrence in Members D and E which are roughly
correlated with Chrons 10 and 9. Contrary to Bishop and Pickford's observation, Hill
et al. (1985) noted that "no equids are known from any of the sites that can be corre-
lated to the Ngorora type section, suggesting an appearance later than 10 Ma." How-
ever, equids dated at between 10.5 Ma and 10.7 Ma have been mentioned in the
Chorora Formation of Ethiopia, associated with some large and small mammals
(Jaeger et al., 1980; Kalb et al., 198Z).
CONCLUSIONS
Review of the available data shows that the synchrony of the Hipparion Datum
throughout the Old World, as suggested by Berggren and van Couvering (1974, 1978),
cannot be demonstrated. On the contrary, when biostratigraphical successions are
well calibrated, some chronologie differences appear in this datum event from one
area to another (figure 3).
Radiometric ages in relation to the Hipparion Datum are very rare and should be
taken into account carefully. In fact, only a few number of Hipparion localities are
associated with lava flows or volcanic ash deposits. No available radiometric dates
can prove that the arrival of the hipparionine equids in the Mediterranean area is older
than 11 Ma.
The magnetostratigraphic calibration of this event has displayed it~ diachrony in

the Siwaliks of Pakistan and in some localities of the Mediterranean area. In the
Siwaliks sequences of the Potwar Plateau, the first Hipparion-bearing horizons in the
Nagri Formation are correlated with the lower part of Chron 9. In the Mediterranean
area sections (Kastellios Hill and Bou Hanifia), Hipparion occurs in Chron 10. Conse-
quently, I proposed an age of about 11.5 Ma (Sen, 1986), which is roughly the age of
502
the Chron 10-Chron 11 boundary, for a possible earliest occurrence of Hipparion in the
Mediterranean area. In any case, a better calibration of this event and its correlation
throughout the Old World require additional data.
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505
THE MAGNETIC STRATIGRAPHY OF THE LATE MIOCENE
SEDIMENTS OF THE CABRIEL BASIN, SPAIN
N.Opdyke
Department of Geology
University of Florida
Gainesville, Florida 3Z611, U.S.A.
P. Mein
Pepartement des Sciences de la Terre
Universite Claude-Bernard-Lyon I
F-696ZZ Villeurbanne Cedex, France
E. Moisaenet
Unite d'Enseignement et de Recherche en Geographie
Universite de Paris I
191 Rue Saint-Jacques
75005 Paris, France
A. Perez-GonAiez
Facultad de Ciencias
Departamento de Geolgia
Universidad de Zaragoza
50009 Zaragoza, Spain
E. Lindsay
Tucson, Arizona 857Zl, U.S.A.
M. Petko
University of Florida
Gainesville, Florida 3Z611, U.S.A.
INTRODUCTION
In recent years, much progress has been made in America and Asia (Pakistan) in
understanding the age range of the Neogene vertebrate faunas and the migration
pattern between these continents. This has been achieved through the application of
magnetic stratigraphy to mammal-bearing sequences of these regions. Unfortunately,
in western Europe the placement of the classic mammal-bearing sequences into the
Magnetic Polarity Time Scale (MPTS) has not progressed so rapidly. One reason for
this lack of progress is the fact that sedimentary exposures in Europe are often poor,
making the application of magnetic stratigraphic techniques difficult.
European Neogene Mammal Chronolov 507

t
N
Sk
6 •
Fig. 1. Locality map of the lower Cabriel Basin, south of Venta del Moro. A =
Cabriel section; B = Venta del Moro section; 1 = Pleistocene traver-
tines; Z = late Neogene continental formations; 3 =Cofrentes volcanic
rocks (middle to late Pliocene); 4 = Limestones, dolomite, and calcar-
enite (Cretaceous and Jurassic); 5 = Clays, gypsum, and dolomite
(middle and late Triassic); 6 = Mammal localities.
THE CABRIEL SECTION: LITHOLOGY AND BIOSTRATIG RAPHY
In an effort to solve this problem, we have selected some of the sedimentary

basins of Spain for magnetostra tigraphy where non-marine sediments are often better
exposed with longer sections {e.g., Jucar and Cabriel valleys, Teruel basin). One
section chosen for, study is in the Cabriel basin, exposed along the road leading from
the Fuente Podrida baths to Los Isidros. The base of the section is represented by
36 m of algal marls and limestones that constitute the upper part of the "Calizas
lacustres de Ia Fuente Podrida" {Riba et al., 197Z; Aguirre et al., 1975). Above, there
is a thick {Z16 m) mainly fluvial sequence. This sequence is formed by sandstones,
conglomerat es and overbank mudstones. It was called the "Miembro de Los Isidros" by
Aguirre et al. {1973), "serie detritique du Cabriel" by Mein et al. {1978), and also "red
polymictic cross bedded conglomerat e, sandstones, mudstones, and concretionar y
carbonate facies" {Mathisen and Morales, 1981).
Four vertebrate fossil assemblages have been recorded in this sedimentary

sequence. They are:
a. Fuente Podrida. This locality, in the Fuente Podrida limestones, was previously
reported by Mein et al. {1978); it contains Parapodemu s barbarae, Occitanomy s
adroveri, and Eliomys truci, and is assigned to the middle Turolian {MN 1Z).
508
b. Balneario. This is a small undescribed site located about 300 m to the east and
roughly 10m stratigraphically below the former; it has provided Parapodemus
lugdunensis, which is a fossil guide to lower Turolian (MN 11).
c. The rich locality of Venta del Moro (Aguirre et al., 1973; Morales, 1981) belongs to
the upper sequence but it is not directly tied to our measured sequence because it
is 7 km to the north. Among the distinctive rodents from this site, we note
Ruscinomys schaubi, Cricetus kormosi, Paraethomys miocaenicus, and
Stephanomys ramblensis. This fauna is characteristic of late Turolian (MN 13).
d. Fuente del Viso. This new locality was discovered last year in the upper part of
the Cabriel Valley on the right bank at Z40 m above the river. Among the char-
acteristic rodents, we note Ruscinomys lasallei, Cricetus barrieri, Paraethomys
anomalus, Stephanomys medius, Apodemus gorafensis, and Blancomys sp. This
fauna represents the latest Miocene in the transition between Miocene and
Pliocene (MN 13/MN 14 boundary) and can be correlated with the Alberca fauna
(Mein et al., 1973).
MAGNETIC TECHNIQUES APPLIED TO THE CABRJEL SEDIMENTS
In the Cabriel valley, samples were collected at 43 sites distributed throughout

the exposure. In order to try and correlate the Venta del Moro site to the road
section, a short section consisting of four sites spanning 10m was sampled through the
Cabriel
Fig. Z. Orthogonal plots of normal and reversely

magnetized samples based on the successive end
points after thermal demagnetization of the
magnetic vectors. Filled squares are vector end
points projected into the horizontal plane, while
the circles are projections on the vertical plane.
509
-··
y p
+oo A
Fig. 3. Lithology and magnetostra tigraphy of the

Cabriel section (left bank of the river) with
V.G.P. latitude plotted against stratigraphic
thickness. Lithology (left-hand column): 1 =
limestones, carbonate silts, clays and marls of
Fuente Podrida Formation. Z, 4 and 6 = sand-
stones, sands, clays, sandy marls, gravels and
petrocalcics of Rio Cabriel Formation. 3 and 5
= carbonate silts and petrocalcics of Rio Cabriel
Formation. Position of sampling sites are indi-
cated by filled circles next to the measured
section. Note that 5 sampled sites in the meas-
ured section did not yield statistically signifi-
cant results. The resulting magnetic polarity
sequence is numbered N1 through R5 in column
A. The shorter Venta del Moro section (column
B) is placed to the right of the measured section,
with possible correlation (to N4 or N5) indi-
cated. Segment of the magnetic polarity time
scale presented to the right.
outcrop. The sites are spaced 5 m apart on average. Three independent ly oriented
samples were taken from each site, utilizing a hand rasp and a brunton compass
(Johnson et al., 1975). The samples were taken preferential ly from fine grained lithol-
ogies such as siltstones, mudstones, or marls, and the sampling of the sandstones was
avoided. The samples were shaped into rough Z em cubes for measuremen t which was
performed on a SCT cryogenic magnetomet er. All samples were incremental ly
thermally demagnetize d in steps to the complete destruction of the remanent magnet-
ization or to temperature s of 680°C. The typical dema~etization program consisted
of 100°C steps to ZOO, Z50, and Z5°C increments to 600 C; above 600°C, the intervals
510
were reduced to 2.0°C. In some cases this resulted in as many as 2.0 discrete demag-
netization steps being performed on some samples.
The resulting data were plotted on orthogonal plots (Ziderveld, 1967, fig. 2.). The
Natural Remanent Magnetization was quite simple, being composed of no more than
two components, as can be seen from figure 2.. In most samples a normal overprint is
present in the direction of the present field. This is more clearly seen in reversely
m~netized samples (figure 2.b) and is removed in most samples by temperatures of
300 c.
A least squares best-fitting line was drawn for linear segments of the orthogonal
plots which were determined by visual inspection (Kirschvinck, 1980). The directions
thus defined were analyzed statistically using Fisher (1953) statistics and converted
into virtual geomagnetic pole positions. These results were then ordered stratigraph-
ically and are presented in figure 3.
BIOSTRATIGRAPHY AND MAGNETOSTRATIGRAPHY: A COMPARISON
It can be seen that the directions of magnetization fall into an ordered sequence
of normal and reversely magnetized sites. These have been designated magnetozones
N1 through N5. Each magnetozone consists of more than one site except for
magnetozone N2. which consists of only a single site and is therefore not as clearly
established. The resulting pattern is distinctive and correlation to the MPTS appears
feasible.
The correlation of this reversal sequence to the MPTS can only be accomplished
through pattern recognition since there are no independently dated units in the
sequence. However, the fauna places the lithostratigraphy clearly in the upper
Miocene. The pattern resembles the reversal sequence of lower Gilbert to Chron 7. In
order to test this hypothesis, a plot of the stratigraphic position against the magnetic
polarity time scale was constructed {Berggren et al., 1985) (figure 4). As seen in
figure 4, the polarity transitions are almost linear, indicating a good fit to the MPTS
for the time from 5 to 7 Ma (Gilbert to Chron 7).
The correlation of these fossiliferous localities in the Cabriel valley with the
magnetic scale brings new information about the age of the faunas.
The Fuente Podrida fauna occurs in the first normal magnetozone (N1) at the
base of the section. Extrapolation of sedimentation rates would place the age of the
Fuenta Podrida fauna at about 7 .2. Ma. Consequently, the boundary between MN 11
and MN 12. should be slightly older than this (about 7.3 Ma) since the Balneario fauna
with Parapodemus lugdunensis occurs only a few meters stratigraphically below the
level of the Fuente Podrida fauna.
The magnetic results obtained at Venta del Moro give the opportunity for evalu-
ating the age of the upper Miocene faunas. In the Venta del Moro section (4 samples),
the upper site was heavily overprinted and weakly magnetized. However, it is clearly
reversed whereas all the other sites are normally magnetized. The fact that a rever-
sal of the earth's field takes place within this short section helps to constrain its
correlation to the road section.
Two alternatives can be envisaged. The first places Venta del Moro in the
N 5/RS transition (boundary Chron 5 - Gilbert); in this case, the fauna should be
5.40 Ma. The second alternative would place the mammals in the N4/R5 transition
and the age of the fauna should be 5. 7 8 Ma.
Two arguments fit well with the second alternative. One is the comparison of
the Venta del Moro fauna with that of Librilla (Thaler et al., 1975). The Librilla fauna
contains roughly the same taxa and lies just above the "lamproitic cinerits" and the
lava flows of the Barqueros volcano which provided the following dates: 7 .00, 6.50,
511
4
X
200m lOOm 0
.,
Rio Cabriel
'"'"" ... "'0 3• .
~s 2
4
1 •••
Fig. 5. Schematic north-south cross-section of the Cabriel valley, with magneto-

zones projected on left bank. 1 = Lacustrine limestones of Fuente Podrida;
Z = Detrital formation of Rio Cabriel; 3 = Lacustrine limestones of Fuente
del Viso; 4 = Pleistocene terraces.
and 6.ZO Ma (Montenat et al., 1975). The second argument is based on the elevation
and faunal composition of the Fuente del Viso mammal assemblage. This fossiliferous
locality is located on the opposite side of the valley exactly at the same elevation as
the NS/RS boundary in the Cabriel section (see figure 5). If the detrital Cabriel
formation is really horizontal, we can place the Fuente del Viso fauna at the NS/R5
boundary. On the other hand, the Fuente del Viso fauna is clearly younger than the
Venta del Moro fauna. Several taxa of the Venta del Morp fauna are represented by
the same genus put with more advanced species in the Fuente del Viso fauna. Fuente
del Viso can also be compared with the Alberca fauna (Mein et al., 1973). In the
second alternative, the Fuente del Viso fauna is placed at the N5/R5 level (5.41 Ma)
and the slightly older Venta del Moro fauna at the N4/R4 level (5.78 Ma).
512
The first alternative for the age of the Venta del Moro fauna (at 5.4 Ma) would
require a lower rate of sedimentation in the area of the Fuente del Viso fauna, or
slight regional dip from the area of the Venta del Moro to the area of the Fuente del
Viso fauna. Either of these scenarios is plausible, which prevents secure assignment at
this time of the Venta del Moro fauna relative to the MPTS.
CONCLUSION
This study shows that it is possible to find long sections favorable for paleomag-
netic investigation in European Neogene deposits. The Cabriel sediments contain 10
magnetozones which correlate with the MPTS from Chron 7 to the Gilbert (7 to
5 Ma). Thus, we have a relatively continuous sequence representing most if not all of
the Turolian land mammal age or stage. In the sampled section, the upper part has
been removed by erosion. This is not a problem on the opposite side of the valley
where the sequence is exposed without apparent gap or discontinuity for 80 m above
the Fuente del Viso site. Consequently, it will be possible to extend and report the
sampling in the area of the Fuente del Viso fauna, which should strengthen age
calibration for the Venta del Moro fauna.
The Cabriel section, because of its length, permits the dating and eventually the
calibration of several MN zones and land mammal ages.
The MN 11/MN 1Z transition is probably near 7.Z0/7.30 Ma.
The middle Turolian faunas (MN 1Z) are about 7 Ma.
The late Turolian assemblages (MN 13) are younger than 5. 7 Ma.
ACKNOWLEDGMENTS
The writers would like to thank the National Geographic Society, NATO, ITGE
(Madrid), CNRS (Paris), Universlte Claude Bernard (Lyon), and Laboratorie de
Geographie physique Pierre Birot (Meudon) for support in the field. We thank also The
Comision Nacional de Geologia (Madrid) and its General Secretary A. Navarro for
authorizing us to work in the Jucar-Cabriel valleys and the Teruel basin. This team is
actively working in the adjacent areas (Jucar valley and Teruel basin) in order to
extend the work reported here in both time and space.
REFERENCES
Aguirre, E., Robles, F., Thaler, L., Lopez, N., Alberdi, M.T., and Fuentes, C., 1973.
Venta del Moro, nueva fauna finimiocena del Moluscos y Vertebrados. Estudios
Geologicos, Madrid, v. XXIX, p. 569-578.
Berggren, W.A., Kent, D.V., and Van Couvering, J.A., 1985. The Neogene. Part Z,
Chronology of the Geological Record," p. Zll-Z60. Blackwell Scientific Publica-
tions, Oxford.
Birot, P. and Sole, Sabaris, L., 1959. La morphologie du SE de l'Espagne. Rev. Geog.
des Pyr. et du Sud-Ouest, v. 30(3), p. 119-Z84.
Brinkmann, R., 1948. Las Cadenas :!3eticas y Celtibericas del Sureste de Espaiia. Pub.
Extranj. Geol. Espaiia, CSIC, Madrid, v. 5, p. 173-Z90.
Fisher, R.A., 1953. Dispersion on a sphere. Proc. R. Soc. London, ser. A, v. Z17, p.
Z95-305.
Instituto Geologico y Minero de Espana, Madrid, 197Z-1978. Geological map of Spain
(1/50000 scale), serie Z, Venta del Moro and Casas lbaiiez.
Jodot, P., 1958. Les faunes de Mollusques continentaux ;reparties dans le Sud-Est de
l'Espagne echelonm!es entre le Miocene superieur et le Quaternaire. Mem. y
Comm. Inst. Geol. Dip. Prov. Barcelona, v. 17, p. 1-134.
513
Johnson, N.M., Opdyke, N.D., and Lindsay, E.H., 1975. Magnetic stratigraphy of Plio-
Pleistocene terrestrial deposits and vertebrate fauna, San Pedro valley,
Arizona. Geological Society of America Bulletin, v. 86, p. 5-1Z.
Kirschvinck, J.L., 1980. The least-squares line and plane and the analysis of paleo-
magnetic data: Geophys. Jour. Roy. Astron. Soc., v. 6Z, p. 699-718.
Mathisen, M. and Morales, J., 1981. Stratigraphy, facies and depositional environ-
ments of the Venta del Moro vertebrate locality, Valencia, Spain. Estudios
Geol~gicos, Madrid, v. 37, p. 199-Z07.
Mein, P., Bizon, G., Bizon, J., and Montenat, C., 1973. Le gisement de la Alberca
(Murcia, Espagne meridionale): correlations avec les formations marines du
Miocene terminal. C. R. Acad. Sc. Paris, t. Z76, ser. D, p. 3077-3080.
Mein, P., Moissenet, E., and True, G., 1978. Les formations continentales du Neogene
superieur des vallees du Jucar et du Cabriel au NE d'Albacete (Espagne),
biostratigraphie et environment. Doc. Lab. Geol. Fac. Sci. Lyon, v. 7Z, p.
99-147.
Montenat, C., Thaler, L., and Van Couvering, J.A., 1975. La faune de Rongeurs de
Librilla, correlations avec les formations marines du Miocene terminal et les
datations radiometriques du volcanisme de Barqueros (Province de Murcia,
Espagne meridionale). C. R. Acad. Sci. Paris, t. Z91, ser. D., p. 519-5ZZ.
Morales, J., 1981. Venta del Moro: su macrofauna de Mamiferos y biostratigraffa
continental del Mioceno mediter~cineo. Tesis Univ. complutense, Madrid, 313 p.
Robles, F., 1974. Levante in "Coloquio internacional sobre biostratigraffa continental
del Neogeno superior y Cuaternario inferior;" Gufa 4.10, Madrid, p. 87-134.
Zidjerveld, J.D.A., 1967. A.C. demagnetization of rocks: Analysis of results, in
Collison, D. W. and Runcorn, S.K. (eds.), "Methods in Paleomagnetism," p.
Z45-Z86. Elsevier, Amsterdam.
514
PRElJMJNARY MAGNETOSTRATIG RAPMC RESULTS
OF SOME NEOGENE MAMMAL LOCALITIES FROM
ANATOLIA truRKEY)
C.G.Laugereia
Paleomagnetic Laboratory
Fort Hoofddijk
Budapestlaan 17
3584 CD Utrecht, The Netherlands
S.Sen
Universite Paris VI
Lab. de Paleontologie des Vertebres et
Paleontologie Humaine
4 Place Jussieu
7 5Z5Z Paris Cedex 05 - Tour Z5, France
M. Siimengen and E. Uuay
Jeoloji Etiidleri
MTA Gen. Md.
Ankara, Turkey
INTRODUCTION
Magnetostratigraphi c research in continental deposits may serve as a useful tool

to correlate widely different mammal assemblages from widely different areas like
Europe and Asia. The role of Anatolia/Turkey in the history of Neogene mammal
dispersal is evident and the present study marks the beginning of a combined magneto-
stratigraphic and mammal biostratigraphic project in this area.
Eight assemblages of rodents have been sampled in the Kayseri-Sivas basin in

Anatolia and faunal analysis shows that they ..range in age from Late Oligocene to
Early Pliocene (Siimengen et al., this volume; Unay and de Bruijn, 1987). Correlation
with western European mammal zones is not straightforward and can generally only be
established tentatively. The present study was aimed at magnetostratigraphi c
sampling in sections containing (some of) the eight mammal localities to get some
more accurate age information on the mammalian assemblages.
GEOLOGY AND SAMPLING
The Kayseri-Sivas basin in Anatolia is a NE-SW trending basin delimited by

metamorphic rocks in the north and the south (figure 1). The basin is divided by an
approximately E-W running thrust fault and there is some uncertainty concerning the
relative ages of the sediments directly north and south of this thrust. The geology of
the basin is described in more detail by Siimengen et al. (op. cit.)
Edited by E. H. lindsay eta/.
(11
~
en
~ Horomi
lignite quarry
..: [Upper [6] limestone

~ U . Miocene
~ Lower~ conglomer ate, sond.Jtone.
l...::_;j siltstone and claystone
..: Mid. Miocene
~ rupperU1I] ~~::!~~eo~~um
.~ Lower (""""""": ~Cind,tone , :tilta~on~,mort
~ L.::::.....:l IJm.etton• al\<l l19M 1le
• Low .- Mid. Miocene

9~sum , Siltstone ..,
~[2'
~ ~ and •ondoton• ·[~..; U.Oiig o- M.Miocene?
m ~ 1 ~~ conglomerate. tandttone
Lo.. 30 eL.'L.J and clo)'11ton•
~ 0
"2 " U.Eocene - OIIgocene? "V
~ ..;- Ant icline "V
Eocene E
·~l11I1 ~~·~:f.d!~!~':"·
~ Synel1ne
C~a~~0°"\y9 ';:r,:. ~ ~~~~~~~u: 11 ~~~~:~~rl11, t ubiditee .JJ- thru't fault
"V"V"V"V_)P+
e rodent locolltl•s v v v/ + + +
Basement Q metamorphic rock• + + +
Fig. 1. Geological map of the Gemerek area (Kayseri-Sivas Basin). The rodent localities are indicated by
numbers: 1 = Yenikoy, Z = Horlak la,b, 3 = Horlak Z, 4 = Gemerek, 5 = Kalekoy, 6 = Karaozii, 7 =
Dendil, 8 = Igdeli. Localities 4, S, and 6 have been sampled for magnetostratigraphic purposes;
locality 8 was not suited for sampling because it was too sandy.
The mammal locality Jgdeli (figure 1) could not be sampled due to unsuitable
lithology (too sandy); some pilot samples have been collected at the Yenikoy locality.
Three sections have been sampled in detail and they contain the Kalekoy, Karaozu,
and Gemerek mammal localities (figure 1). It was tried to sample every section over
as long and continuous stratigraphic range as possible - depending on suitable lithol-
ogy and tectonics --in order to get a maximum number of polarity zones to facilitate
correlation with the polarity time scale (cf. Langereis et al., 1984). Care was taken to
sample in fresh sediment - if that was possible - by removing the weathered
surface. Jn all sections, inter-level distances were calculated according to Langereis
and Meulenkamp (1979).
The Kalekoy and Karaozii sections are of late Vallesian age (Sumengen et al., op.
citJ and consist of (mainly chocolate-brown) clays, silts, and sands. The KalekCiy
section is somewhat older than the Karaozii section but the age difference seems
minimal since the sections are situated directly above and some 50-100 m stratigraph-
ically above the same prominent gypsum layer. The Kalekoy section contains thick
layers of sand which are not suitable for taking paleomagnetic samples, whereas the
Kara8zii section consists mainly of silts and clays. However, we note here the great
lithological similarities in the sediments of these two sections. The bedding planes
have a strike and dip of 195°S, 45-55°W or Z60°W, 45°N. Cores were mostly drilled in
the field with an electric drill (and a generator as power supply) using water as a
coolant, but at a number of levels handsamples were taken because drilling with water
dissolved too much of the core. From the handsamples we tried to drill cores in the
laboratory using compressed air. Jn the Kalekoy section Z1 levels were sampled over a
stratigraphic range of 60 m; at 3 levels no cores could be obtained. Jn the Karaozii
section 31 levels were sampled over a stratigraphic range of 5Z m; at 5 levels no cores
could be obtained.
The Gemerek section is roughly of Aragonian age (Sumengen et al., op. cit.) and
shows a somewhat irregular cyclicity of sands, silts, marls/clays, limestones, and
lignites; the cycles are often incomplete, restricted to only sand~ silts, and some
clays. The bedding plane has a strike and dip of ZZO-Z40°S, 60-80 W. Jn the lower
part (0-50 m level) the sediment is very sandy and silty, in the middle part (50-1ZO m
level) there are intercalated but relatively thin (1 m) limestones and lignites, and in
the upper part of the section (lZ0-170 m level) the limestones and lignites can reach a
considerable thickness (3-5 m). Oriented handsamples were taken at 6Z levels over a
stratigraphic range of 170 m. The interval of the sediment sequence between the
170 m level and an overlying basalt is not exposed, resulting in a stratigraphic gap of
approximately 80 m. Above this gap, the section contains an intercalated basalt flow
which has a K/Ar date of 14.9 ± 0.9 Ma. From the handsamples, cores were drilled at
the paleomagnetic laboratory in Fort Hoofddijk, after embedding the samples in
plaster of Paris because of the fragility of the sediment.
LABORATORY TREATMENT
The natural remanent magnetization (NRM) was measured on a ZG Enterprises

cryogenic magnetometer. The total NRM intensities are generally very high, typically
10.0-80.0 mA/m for Kalekoy and 4.0-50.0 mA/m for Karaozii. The total NRM intensi-
ties are intermediate in the Gemerek section, 0.5-5.0 mA/m. Jn general two speci-
mens per sampling level have been subjected to progressive thermal demagnetization,
using small temperature increments, up to a maximum of 650°C.
Kaleltliy and Karai:SzU
Thermal demagnetization of samples from the Kalekoy and KaraozU sections

reveals that the total NRM is generally composed of three components (figures Z and
3). A small viscous and randomly directed component is removed at temperatures of
100°C and can be regarded as being laboratory induced. An-- often relatively large -
secondary component is usually removed at temperatures of ZOO-Z50°C. This second-
ary component has a present-day direction (before bedding tilt correction) and is
517
I!L_u TK 10 1 TK 11 2A
up/W up/W
70.0 21.0
up/W
100"C 26.0
200 100"C
200
100 'C
650 N
10.0
300 N
20.0
\
-~
~--
N
-eo.o -10.0
Fig. Z. Thermal demagnetization diagrams of specimens from the Kalekey

section. Open (solid) circles denote orthogonal projection (after
bedding tilt correlation)
.
on the vertical (horizontal) plane; numbers
0
refer to temperatures in C. Often, a very large secondary component
is present, overlapping the ChRM to a large extent. Nevertheless,
ChRM directions can be interpreted as being originally of reversed
polarity.
therefore of recent origin and most probably caused by weathering of the sediment.
Often a reversed characteristic remanent magnetization (ChRM) component is finally
removed at temperatures higher than 330-360°C, up to a maximum temperature of
610-650°C. Between the secondary and characteristic component there appears to be
an overlap in their blockin§ temperature spectra. This overlap seems to range
between Z50°C and 300-330 C if the secondary component is relatively small (e.g.,
TK Z 1 in figure Z; TK 71 1 in figure 3), and between Z50°C and 390°C or even as high
as 4Z0°C if the secondary component is large compared to the ChRM component (e.g.,
TK 11 ZA in figure z; TK 56 1 and TK 57 ZA in figure 3). This seemingly variable
overlap in blocking temperature spectra is due to a larger ratio of secondary and
characteristic component (Langereis, 1984), e.g., because of a larger secondary
component resulting from increased weathering. In most cases a "window" of non..,.
overlap in the ChRM blocking temperature spectrum remains, which makes it possible
to recognize the (reversed) ChRM from the overprint. In some cases, however, the
intensity of the secondary component is too large to determine reliably the direction
of the ChRM (e.g., TK 57 ZA in figure 3). In such a case, only a direction intermediate
between secondary component and ChRM is determined and plotted (figure 4).
The ChRM and intermediate directions of both the Kalekoy and Karaozii sections
(figure 4) therefore sometimes show positive inclinations or declinations deviating
from south due to extensive overprinting of this secondary component. Nevertheless,
all directions determined from both sections are interpreted to be indicative of
original remanent magnetizations with a reversed polarity.
Gemuelt
Thermal demagnetization of samples from the Gemerek section shows in general

very similar results with respect to the different components (figures 5 and 6). In this
section, normal polarity ChRM components are also seen (figure 5). This normal
ChRM component can be easily distinguished from the secondary component due to
the considerable tilting of the bedding plane (60-80~. Furthermore, the recognition of
true normal polarity is facilitated by a large counterclockwise rotation of ca. 50-60°
518
TK 77 1
up/W
up/W 15.0
15.0 300
500
TK 56 1
up/W
N
~
~-roo•c
N
~0
•
-11.0 500
-11.0
TK 57 2A
up/W
11.0
-2.0
Fig. 3. Thermal demagnetization diagrams of specimens from the

Karaozii section. See also caption of figure z.
of the section, and therefore the clear directional difference between normal polarity
(310~ and normal overprint (360~. At the same time, this indicates that the normal
polarity ChRM is most probably of primary origin. The large counterclockwise rota-
tion is thought to be due to local tectonics, considering the local direction of the anti-
cline axis SE of the section (figure 1).
A secondary component is removed between 100° and ZOO-ZS0°C and clearly has
a present-day direction before bedding tilt correction; the mean direction of this
secondary component has a declination of 0.5° and an inclination of 55.7°, which is
close to the geocentric axial dipole field inclination (58~.
The ChRM components in the Gemerek sections are generally totally removed at
390-4Z0°C, which is at considerably lower temperatures than in the previous two
sections. This may indicate that the carrier of the ChRM is titanomagnetite (Dankers,
1978; Hartstra, 1982). The basalt flow in the top of the section suggests that volcanic
activities have taken place approximately at the time of deposition, which would
indeed promote the occurrence of Ti-rich minerals. Similar to the Kalek()y and
Karaozii sections, the ChRM directions often cannot be distinguished reliably and
intermediate directions are found (figure 6), but these are interpreted to indicate
original remanent magnetizations of both normal and reversed polarity.
519
ChRM - directions KALEKOY ChRM - d ir ec II ons KARAOZU
N N
Fig. 4. Equal area projection of the ChRM and intermediate directions of the
Kalekoy and Karaozu $eCtions. Open (solid) circles denote projection on
the upper (lower) hemisphere. The ChRM directions are sometimes not
well established; the resulting intermediate directions show positive
inclinations and/or declinations deviating from south, due to a large
secondary magnetization overprint.
TG 12
up/W up/W
100 oc tOO 100 "C
200
200
N
2.50
N
-0.50
2.50
Fig. 5. Thermal demagnetization diagrams of specimens from the Gemerek section,

showing normal polarity ChRM directions, which can be easily distinguished
from the secondary overprint due to considerable tilting of the bedding
plane and (local) rotation of the section. See also caption of figure 2..
620
TG 42 1A
up/W
0.25
1oooc
TG 44 1A
up/W
0.40
N
0.25
300
N
390
-o.10
-o.111
TG 45 1A TG 5U
up/W up/W
0.4 1.00
200
N
390
-0.25
300
390 N
1.50
-o.4 -0.50
Fig. 6. Thermal demagnetization diagrams of specimens from the Gemerek

section, showing reversed polarity ChRM directions. Note that despite
the large secondary overprint, the characteristics of the demagnetization
diagrams are entirely different from those of figure 5.
MAGNETOSTRATIGRAPHY
KaieJr.c;y and Karaazu

The mammal fauna from both localities m::e of late Vallesian age. According to
Berggren et al. (1985) the top of the Vallesian is ca. 9.0 Ma and the base is ca.
1Z.5 Ma. Both the Kalekoy and the Karaozii sections yield only reversed ChRM direc-
tions and hence consist of only one reversed polarity zone, which would imply that the
sections are either younger than the top of anomaly 5 (8.9Z Ma) or older than the base
of anomaly 5 (10.4Z Ma; see figure 8). If both sections are younger than 8.9Z Ma, the
Vallesian/Turolian boundary would be younger than 8.9Z Ma. The other possibility,
that both sections are older than 10.4Z Ma, is equally possible from a magnetostrati-
521
180
I
0 0
I 0
0 0 00 0
0 0
I
0 01
.
•
~
0
0
-; 120 120
E co
(][)
0 0 0 0
I
•> 'o
100o 0
I
I
• 0
Cf,o 0 0
(II)' 0
CD
80
:;:: I 0
.
0 10 0
~ (][) I .!IP 0 o
I Oo 8f"
40
20 20 ' CDo
0 0 0 I 0
0
0
I I 0~-T~~-T~~-T~
80 18 270 .1110 -10 -10 -.110 0 .110 10 80
declination inclination
Fig. 7. Declination and inclination of ChRM and intermediate directions
versus stratigraphic level in the Gemerek section. White (black)
denotes reversed (normal) polarity. A counterclockwise rotation of ca.
50-60° is indicated by arrows.
graphic point of view; but since the mammal fauna appears to be quite close to the
Turolian stage or age, this solution is less appealing. Correlating the sections to the
reversed sub,chron directly above anomaly 5 (8.80-8.9Z Ma) would in both sections
imply a sedimentation rate of more than 40-50 cm/ka. In the case of a correlation
with the reversed subchron between 8.50-8.71 Ma this sedimentation rate will
decrease to more than Z5-30 cm/ka. Such sedimentation rates seem reasonable for
continental deposits (Sen et al., 1986; Sen and Valet, 1986; Sen, 1988).
One more possibility needs to be considered; the existence of very short reversed
subchrons in anomaly 5 has been suggested (see LaBrecque et al., 1977), but they are
on the order of Z0-30 ka. This would mean that the sedimentation rate must be more
than Z00-300 cm/ka. This is very high compared to other continental deposits (Sen,
1988) and therefore less likely. In the Kalek(iy section there are some very thick
sandy channels, which may be regarded as being deposited instantaneously. If thick-
ness of these channels were subtracted from the total thickness, it would decrease
approximately to half its former value and consequently sedimentation rates would be
less by a factor of two. On the other hand, in KaraozU no similar channels have been
observed, and the above argumentation would apply to Kalekoy as well because of the
time equivalence.
522
8
10
11
..
...
12
..,a
>-
1::
0
13
=
E
t
~ 1-4 14.9 ± 0.7 Ma
..,
Cl
<(
15 II
>12 >12
16
23 14
17
>50 >30
18
19
20
Fig. 8. Possible correlations of the Kalekoy I
Karaozii and the Gemerek sections to the
polarity time scale based on sea-floor
anomalies (Berggren et al., 1985). Pre-
ferred correlations are indicated. For the
Gemerek section the preferred correla-
tion shows two possibilities (I and II); the
corresponding inferred sedimentation
rates in cm/ka are listed for the three
polarity zones.
Gemerek
The Gemerek section consists of three polarity zones (figure 7); it starts at the
bottom with a small reversed zone (of ca. 20 m), followed by a normal zone (of ca.
70 m), and to the top of the section by a reversed zone (of ca. 75 m). On top of that
there is a gap (ca. 80 m) followed by a basalt.
523
On the basis of the K/Ar date of the basalt in the (unsampled) top part of the
section, the section is older than 14.9 ± 0.7 Ma. Using the Berggren et al. (1985)
polarity time scale based on sea-floor anomalies, there are several possibilities to
correlate the magnetostratigraphic sequence of the Gemerek section (see also figure
8), Some reasonable possibilities are to correlate the normal polarity zone with the
normal chrons (a) 14.87-14.96 Ma, (b) 15.13-15.27 Ma, (cl) 16.22-16.52 Ma, or
(c2) 16.22-16.73 Ma. The latter possibility (c2) would imply that we missed the very
short (40 ka) subchron from 16.52-16.56 Ma. The various possibilities imply the
following (minimum) sedimentation rates (in cm/ka) and ages for the Gemerek
mammal fauna.
.!. B. .£!. c2
reversed (75 m) >35.0 >44.0 >12.0 > 12.0

normal (70 m) 78.0 50.0 23.0 14.0
reversed (20 m) >n.o >2.0 >50.0 >30.0
age of fauna (Ma) 14.9 15.2 16.4 16.5
It is not possible to establish a firm magnetostratigraphic correlation on the

basis of the current polarity sequence, but considering the lithology of the Gemerek
section, correlation c1 is most appealing. The lower part consists of sand and silts
(>50 cm/kal, the middle part contains thin limestones (23 cm/ka), and the top part
contains thick limestones (>12 cm/kal. Since it is very likely that limestones (and
lignites) have a much lower sedimentation rate than sands and silts, we consider cl the
most reliable though tentative corelation, yielding an age of 16.4 Ma for the Gemerek
mammal fauna. The correlation is, however, far from definite. We have, therefore,
extended the Gemerek section further downward this summer, and also we have
sampled the basalt flow and the limestones on top of this flow.
CONCLUSIONS
We have sampled three sections containing the Kalekoy, Karaozii, and Gemerek
mammal faunas (Si.imengen et al., this volume). Our conclusions are tentative and are
as follows:
1. The magnetostratigraphy of Kalekoy and Karaozi.i sections corroborates the

assumed synchrony of the sections since both are entirely of reversed polarity.
The sections are probably younger than the top of anomaly 5 (8.92 Ma).
2. The magnetostratigraphy of the Gemerek section, plus the age constraint of

14.9 ± 0.7 Ma, yields several possible correlations. On the basis of the implied
sedimentation rates and the lithology of the section, a correlation yielding an age
of 16.4 Ma for the Gemerek mammal fauna is at present preferred.
ACKNOWLEDGMENTS
Hans de Bruijn was very helpful, both in the field as well as in giving helpful
comments. Piet-Jan Verplak did most of the laboratory measurements.
REFERENCES
Berggren, W.A., Kent, D.V., Flynn, J.J., and Van Couvering, J.A., 1985. Cenozoic
Dankers, P, H. M., 197 8, Magnetic properties of dispersed natural iron-oxides of known
grain-size. Ph.D. thesis, University of Utrecht.
Hartstra, R.L., 1982. Some rock magnetic properties for natural iron-titanium
oxides. Ph.D. thesis, University of Utrecht.
524
LaBrecque, J.L., Kent, D. V., and Cande, S.C., 1977. Revised magnetic polarity time
scale for Late Cretaceous and Cenozoic time. Geology, v. 5, p. 330-335.
Langereis, C.G., 1984. Late Miocene magnetostratigraphy in the Mediterranean.
(Ph.D. thesis, University of Utrecht), Geologica Ultraiectina, 32, 180 p.
Langereis, C.G. and Meulenkamp, J.E., 1979. Comparison of two methods used to
measure lithostratigraphical intervals in sections Potamidha 1 and 2. Utrecht
Micropal. Bull., v. 21, p. 22-26.
Langereis, C.G., Zachariasse, W.J., and Zijderveld, J.D.A., 1984. Late Miocene
magnetobiostratigraphy of Crete. Marine Micropal., v. 8, p. 261-281.
Sen, S., 1988. Magnetostratigraphie et taux de sedimentation: quelques donnees sur
les depots fluviatiles, lacustres et marins du Neogene mediterraneen. Bull. Soc.
geol. France, (8), t. IV, no. 1, p. 161-166.
Sen, S. and Valet, J.P., 1986. Magnetostratigraphy of Late Miocene continental
deposits in Samos, Greece. Earth and Planetary Science Letters, v. 80, p.
167-174.
Sen, S., Valet, J.P., and Ioakim, C., 1986. Magnetostratigraphy and biostratigraphy of
Siimengen, M., Unay, E., Sarac, G., Bruijn, H. de, Terlemez, I., and Giirbuz, M., this
.. volume. New Neogene rodent assemblages from Anatolia (Turkey).
Unay, E. and Bruijn, H. de, 1987. Middle Oligocene to Early Miocene rodent assem-
blages from Turkey, a preliminary report. Munchner Geowiss. Abh. (A), v. 10, p.
203-210.
625
THE ClUNESE NEOGENE MAMMALIAN BIOCHRONOLOGY-
rrs CORRELATION WITH THE EUROPEAN NEOGENE

MAMMALIAN ZONATION
Zbanxiang Qiu
Institute of Vertebrate Paleontology

and Paleoanthropology
Academia Sinica
Beijing, China
HISTORICAL REVIEW AND RECENT PROGRESS
In contrast with Europe, the East Asian mainland is mainly characterized by

Cenozoic continental deposits. Its coastline has been little altered at least since the
Neogene, resulting in limited deposition of marine sediments on the mainland.
Continued subsidence of the coast area during most of the Cenozoic Era has deeply
buried the few marine deposits. On the other hand, the strong uplift of the southern
and central parts of Asia, enormously intensified since the beginning of the Neogene
as evidenced by the gigantic latitudinal mountain systems of the Himalayas, Tienshan,
etc., has provided an immense amount of terrigenous material to the continental
basins. Therefore, Neogene deposits, often thousands of meters thick and richly
fossiliferous, are widely distributed in East Asia, especially in North China. South
China, being principally a zone of denudation, is devoid of significant Neogene
deposits, except for some small patches in Yunnan Province and Guangxi Zhuang
Autonomous Region. For this reason the Neogene stratigraphy of China has been
almost exclusively based on the continental deposits, and the Neogene geochronology
has relied on mammalian paleontology.
Historical ReView
Leaving out the disjunct records of the single pioneer explorers, the history of
Neogene research in China shows an unusual bias toward the discovery and elaboration
of the fossil mammals, often with little geological documentation. By the middle of
the present century several hundred localities of Neogene mammal fossils had been
found and a number of monographs devoted to description of these fossils had been
accomplished, but only a few papers were devoted to the Neogene stratigraphy, mostly
of preliminary reconnaissance character.
The first effort to subdivide the Neogene of China was made by J .G.
Andersson. In his "Essays on the Cenozoic of North China" published in 19Z3,
Andersson recognized two Neogene units: the Hipparion clays of Shanxi, Henan,
Shaanxi, and Gansu Provinces of "upper Miocene or lower Pliocene" age and cave-
deposit of Chou-Kou-Tien of "upper Pliocene" age. The most important localities of
Hipparion fauna listed by Andersson were Chi Chia Kou in Baode County, Nan Sha Wa
in Hequ County of Shanxi Province, Wu Lan Kou in Fugu County of Shaanxi Province,
Hsin Chia Kou and Chao Chia Chain Qingyang County of Gansu Province, Ertemte of
Nei Mongol and some localities in Xin-an and Luoyang of Henan Province. Andersson
did not come to a final conclusion as to the age of the Ertemte localities because the
fossils he had collected were then still under study. However, he suggested that they
might be a little younger than the other localities with Hipparion. Andersson also
introduced the San Men Series to the Cenozoic stratigraphy of China. The sub-loess
yellow sands and gravels rich in freshwater bivalves along the middle reach of the
Yellow River were first noticed by V.K. Ting. Andersson believed the San Men Series
must have been closely related to loess, hence he placed it in the Pleistocene, younger
than the Chou Kou Tien fissure-filling deposits, which he thought to be of Pliocene
age. As regards the age assignment of the Chou Kou Tien cave deposit, Andersson's
mistake became apparent soon after the Peking man fossils had been found in this
cave depo§it. As a result, only the Hipparion clays of "upper Miocene or lower
Pliocene" age were left in Andersson's Neogene of China.
In 19Z4 M. Schlosser published his monograph on the Ertemte vertebrate fossils.

He gave a complete description of the fossils and thorough discussion on the affinity
of the fauna. Schlosser was of the opinion that the Ertemte fauna was closer in affin-
ity to the Roussillon fauna than to those mammalian assemblages thought to represent
the Pontian of Eurasia. Unfortunately, all the other students who were invited to
study the fossils of the large Lagrelius Collection of the Chinese Neogene mammals
confined their efforts to paleontological work and made little effort to clarify the
stratigraphic position of these remains.
From 19Z1 through 1930 the 3rd Central Asiatic Expedition organized by the
American Museum of Natural History had been conducting an unprecedentedly large-
scale exploration and excavation of vertebrate fossils in the Mongolian Gobi area.
One of the sensational discoveries was that of the shovel-tusked Platybelodon at
Tung-gur in 19Z8-30. Instead of Hipparion, Anchitherium was found there, so it soon
became clear that the Tung-gur fauna represented a new pre-Hipparion fauna in
China. Otherwise, the American expedition made little contribution to the Chinese
Neogene stratigraphy.
Through more than thirty years of persistent work, P. Teilhard de Chardin

undoubtedly played a key role in establishing the framework of the Chinese Neogene
biostratigraphy, often in collaboration with his Chinese colleague, C.C. Young. It was
he who first in 19ZZ unequivocally linked the Chinese Hipparion fauna with the
European term "Pontian," which became, unfortunately, too deeply rooted in Chinese
Neogene terminology.
Teilhard de Chardin also played a decisive role in transferring the usage of

Miocene and Pliocene from the European concept to the American one. During the
19Z0s most of the students who worked on Neogene problems of China adopted the
West European practice of including the Pontian Stage in Miocene. This point of view
had been held by Teilhard de Chardin until 19Z9 when he and C.C. Young suddenly
placed Pontian in the Pliocene in an appended table to a report on the excavation at
Chou Kou Tien (actually published in 1930). Since then, until the end of the 1970s, the
Chinese Hipparion fauna had consistently been considered as belonging to lower
Pliocene. Teilhard de Chardin did not clearly explain why such a change of his
attitude occurred until 1934, when he published a paper jointly with R.A. Stirton
specially devoted to the Miocene-Pliocene boundary problem. Teilhard de Chardin was
of the opinion that a complex scale reflecting both marine and continental division
would be the best way to suppress the discrepancies between the two schools of
stratigraphers. As far as the continental division of the European Neogene was
concerned, he was in full agreement with the suggestion put forward by W.D. Matthew
that the base of the Pliocene should be marked by the immigration event of Hipparion
from North America to Eurasia.
Probably Teilhard de Chardin's most important contribution to the Chinese

Neogene was to fill the stratigraphic gap between the Hipparion red clay and the
loess. We owe our knowledge of the famous Nihewan mammalian fauna to his effort.
The importance of the fossiliferous deposits in the Nihewan area was first recognized
by G.B. Barbour and Teilhard de Chardin (19Z4). Later, mainly through E. Licent's
efforts in the next three years, a rich mammalian fauna was collected. The fauna was
528
of significance on two counts. First, based on freshwater bivalves, Teilhard de
Chardin concluded that the Nihewan deposits were contemporary with Andersson's San
Men Series. Second, the fauna was believed to represent an intermediate one linking
the Hipparion fauna, on the one hand, with the Coelodonta antiquitatis fauna of the
loess on the other. He believed there were three major Cenozoic divisions in China:
the Hipparion fauna, the intermediate Hipparion-Equus fauna, and the Coelodonta
antiguitatis-Bos primigenius fauna. As regard the geological age of the Nihewan
fauna, Teilhard de Chardin considered it as belonging to late Pliocene-early Pleisto-
cene. Before the large-scale excavation at Chou Kou Tien was conducted, Teilhard de
Chardin had included Chou Kou Tien in his Sanmenian in a broad sense. However, as
soon as he made acquaintance with the rich material obtained during the first excava-
tion campaign in 192 7-29, he at once placed Chou Kou Tien into the Pleistocene,
above the Sanmenian.
Teilhard de Chardin believed that the hiatus between the Hipparion fauna and
that of Nihewan was larger than that between the Nihewan and the loess. That is why
he repeatedly returned to the topic of the Sanmenian and tried hard to subdivide it
further. In 1930 he and C.C. Young recognized the "reddish clay" as a lithostrati-
graphic unit lying between the Hipparion red clay and the loess. They divided the
reddish clay into three zones based mainly on the succession of a particular group of
cricetid rodents, the Myospalacinae. Thus Zone A was characterized by M. omegodon,
Zone B was characterized by M. tingi, and Zone C by M. fontanieri. Teilhard de
Chardin synchronized Zone B with Nihewan and C with Chou Kou Tien. Zone A was
thought to be a new level, probably belonging to middle or upper Pliocene, roughly
comparable with Ertemte and the white beds of Dalai Nor, but not with Jingle, which
was then still considered to belong to the Hipparion red clay, although Teilhard was
quite aware of the higher evolutionary level of the forms of Jingle fauna than those of
the typical Hipparion fauna.
In the last ten years of his stay in China, Teilhard de Chardin devoted himself
mainly to the monographic study of the rich material collected from Yushe, Shanxi, by
Licent in 1934-1937. He adopted Licent's tripartite subdivision of the Yushe series.
Yushe Zone I corresponded to the Hipparion fauna of the early Pliocene. Zone ll was
characterized by many new forms like advanced deer, antelopes, etc. Teilhard de
Chardin attributed it to middle Pliocene, and correlated it with Jingle and Ertemte.
Yushe Zone m was correlated with Nihewan, and considered late Pliocene in age,
mainly based on its close resemblance with the Villafranchian fauna in Europe.
Teilhard de Chardin still held this viewpoint when he left China in 1945.
As far as the deposits earlier than Hipparion red clay are concerned, Teilhard de
Chardin and Young could find only two of them during the 1930s: Shanwang of
Shandong Province and Cixian of Hebei Province. From the former locality they
described a new genus of rhinoceros, Plesiaceratherium, several genera of primitive
deer, a carnivore, Amphicyon, etc. The fauna was assigned to late Miocene, compara-
ble with Tung-gur. The Cixian fauna was very poorly represented, and was mentioned
only in an appendix to the description of the Shanwang fauna.
In the Lagrelius Collection there were also some fossils that came from deposits
evidently older than the Hipparion red clay. H.S. Pearson (1928) first described some
teeth and bones of Listriodon gigas from Xianshuihe, Gansu. Pearson regarded the
fossiliferous deposits as belonging to the same age as Tung-gur. In 1936 C.C. Young
and M.N. Bien revisited the locality and added some new forms to the fauna. A.T.
Hopwood described some proboscidean teeth discovered from around Xining of Qinghai
Province in 1935. This locality was roughly regarded as belonging to the Miocene. B.
Bohlin also made some contributions to the Neogene of China by adding some new
localities. He published his monograph on the Qaidam (Tsaidam) fauna (Bohlin, 1937).
The presence of Hipparion tooth fragments in association with a series of new forms,
often more primitive than their counterparts in the typical Hipparion fauna, made
Bohlin believe that the Qaidam fauna was older than the Hipparion fauna. While
describing the rich Oligocene mammalian fauna from the Taben-buluk area, Bohlin
(1946) mentioned a Gomphotherium-Sayimys-Kansupithecus level; he compared it
directly with Tung-gur and thought them contemporaneous.
529
This was the general framework gradually developed and formulated during the
period from the 19ZOs through 1940s. Mainly because of the war, and some other
political reasons, the Neogene study in China then remained rather stagnant until the
end of the 1970s. Among the odd discoveries of that period, the most important were,
for example, the lower jaw fragment of Anchitherium from Fangshan near Nanjing
(Chow M.c. and Hu C.k., 1956), Dryopithecus from a coal mine of Kaiyuan, Yunnan
(Woo J.k., 1957), the listriodont jaws from Lantian area near Xi-an (Liu T.s. and Li
Y.q., 1963), and a small fauna from Cixian (Chen G.f. and Wu W.y., 1976). Their
geological age was only roughly determined as Miocene, without accurate correlation
(see table 1).
Recent Progress
Probably 1978 might be regarded as a turning point in the history of the Chinese
Neogene study. From that year on, it has been possible to publish the results of the
Neogene work done shortly before or during the so-called culture revolution. Also in
that year, a Neogene group in the Institute of Vertebrate Paleontology and Paleo-
anthropology, Academia Sinica, was formed, with the aim to resume the work in a
more systematic way than before. Since then, considerable progress has been made.
Intensive survey of the Cenozoic deposits in the Lantian area had been con-
ducted for three successive years, from 1963 to 1965. As a result of the survey, some
new levels of Neogene deposits were found. The results were published in 1978 (Zhou
M.z., 197 8). Lengshuigou Formation was believed to belong to the middle Miocene,
stratigraphically lower than the Tung-gur Formation. From it some "Hispanotherium"
and "Listriodon", etc. were found. The Babe Formation was considered stratigraphi-
cally lower than the typical Hipparion red clay. From the Babe Formation two new
Hipparion species, a giant "Percrocuta", and other taxa were found.
Xu Q.h. et al. (1978) reported the discovery of a Ramapithecus jaw in a late

Miocene lignite deposit of Lufeng, Yunnan. The excavation actually started in 1976,
and has been carried on intermittently during the last ten years. A number of papers
have appeared since 1978, but the bulk of the material is still under preparation and
study. The age of the fauna has been estimated as about 7 to 8 Ma, according to the
evolutionary level of Brachyrhizomys. This is the only mammalian fauna of late
Miocene age ever found in south China.
In 1980 the results of the Tibetan expedition organized by the Academy of

Sciences {1973-1976) in the field of vertebrate paleontology were published. The
discovery of a Hipparion fauna at two localities, Bulong and Guizhong, was the first
Neogene mammal record from the Tibetan plateau.
In a series of papers Li C.k. and Qiu Z.d. (1980) and Qiu Z.d. et al. (1981)
reported new finds in the Xining Basin of Qinghai Province. The predominantly
micromammalian fauna of the Xiejia Formation was unquestionably of prime impor-
tance. It represented the first authentic record of early Miocene mammals in China.
In 1980 a group of Chinese and West German paleontologists procured a very rich
collection of micromammals in the classic area of Ertemte by the screen-washing
technique. The preliminary report appeared in 1983. Since then, several mammalian
groups of the collection have been published, and a large part of the material is still
under study. The fauna has been correlated with MN zone 13 of the European
mammalian zonation.
A wide range of prospecting and excavation of Neogene mammals has been

conducted lately in North China. As a result, faunas of considerable importance have
been found. For example, exquisite specimens of old and new taxa were unearthed
from the diatomite quarry of Shanwang; from Sihong, Jiangsu Province, a great
number of isolated teeth of various taxa supposedly of an age earlier than that of
Shanwang were collected; many skulls and jaws of both large and small mammals of a
middle Miocene .fauna were purchased from the Tongxin area, Ningxia; very interest-
530
Table 1. Subdivisions of the Neogene terrestrial deposits and faunas in China prior to 1980.
Andersson, 1923 Schlosser, Zdan- Teilhard de Ghardin et Bohlin, 1946 Zhou M. z., Chiu z. x, et Epochs
sky et al., Young, 1930 - 1945 1978 al., 1979 Rogl et Stei-
1924-1930 ninger, 1983
Pleis Choukoutien Pleis. Choukoutien c .J P1eist.
.J . - - 1.8 - -
Ssn Men Nihowan Nihowan .. 7 'Gl Ssmlen
'G"'l (=Ssmlen) t:? li! .,
li! Series (=Ssmlen)
s::" Yushe III 8 1-
Choukoutien .<:1
.,s::
" A
"0 ? "0
Ching1o
j"' Ching1o .....
.... L--L..-
., ..... Yushe II ....
s:: 0.. 0..
., ., ., ~5.4-
I-- s::
Ertemte s::
"
0 "' ()
"
..... "
0
..... Paote Paote Lantian 0 Paote
Hipparion .....
0.. Red Clay .....
..... .....
I
0..
0 Baho 0.. Baho
..... "s:: Tsaidam
:&:
"
0" Tung-gur Hsishui (Ta- Koujiacun 'l\lnggur .,
..... Shanwang ben IW.uk)
X Hsianshuiho <I
etc,
., .,
"
<I
Lengshuigou
<I ()
" ()
"' Shanwang 0
0
0 0
..... .....
.....
X X X
Xiejia
Ul
w
ing fossils of early, middle, and late Miocene were obtained from a wide area of the
middle part of the Gansu Province.
Recently, in 1986 and 1987, we organized a large scale excavation at the classic
locality Tung-gur. From one "graveyard" no less than ZO individuals of Platybelodon
were found, counted by skulls and jaws. Altogether, more than 100 large boxes were
sent back to the laboratory. Since 1987, a joint Sino-American project incorporating
paleomagnetic dating with conventional geologic-paleontologic work has been carried
on in the Yushe Basin. Just at the beginning of 1988 we succeeded in purchasing about
ZOO chilothere skulls and about 40 skulls of other genera common in the Hipparion
fauna from Fugu, a county on the west bank of the Yellow River opposite the famous
fossiliferous area in Baode County. All these materials have not been prepared.
Based on the available material, Li C.k. et al. (1984) gave a preliminary

summary of the Chinese Neogene mammalian faunas in tabular form. However, the
rapid progress in Neogene study of China in the last years makes it possible to improve
considerably the framework proposed by Li et al. (see table Z).
NEOGENE MAMMALIAN FAUNAL SUCCESSION OF CHINA
The last ZO years have witnessed great progress in the field of Neogene stratig-
raphy, especially in Europe ap.d North America. Through multidisciplinary and coordi-
nated efforts of a large number of specialists, refined and reciprocally calibrated
biostratigraphic and/or chronostratigraphic tables and correlation charts have been
proposed and recommended for west Europe, middle and east Paratethys, and North
America, respectively. To propose such a comparable table for the Neogene of China
seems now a little premature. As a matter of fact, except for some well-known
localities, like Baode, Tung-gur, and Nihewan, which have no less than 50 years'
history of investigation, what we now possess are mostly isolated and only preliminar-
ily studied assemblages of fossil mammals. For the majority of the assemblages to be
discussed below, and even the well-known faunas, either systematic collecting or
geological work or both are badly needed. This renders it absolutely necessary to
refer as much as possible to the progress achieved recently in Europe and North
America in establishing a correct sequence of the Chinese Neogene mammalian
faunas. A priori, we have taken the European faunal succession as the main standard
for comparison because now Asia and Europe constitute the main part of a single
zoogeographic region, the Palaearctic region. The paleozoogeographic problems will
be discussed in detail in the section that follows, but it is safe to say that during the
Neogene both Asia and Europe had more faunal elements in common with each other
than with other continents.
Lanzhou Fauna
A local fauna discovered in 1986 south of the Lanzhou railway station is com-
prised of five forms: Metexallerix gaolanshanensis, Tataromys suni, Tataromys sp.,
Leptotataromys gracilidens, and Tsaganomys cf. altaicus (Qiu z.x. and Gu Z.g.,
1988). The fossils were found in gypsiferous red clay, which has been tentatively
correlated to the top of the second formation (Yehucheng Formation) of a thick rock
series well developed on the north bank of the Yellow River. In his last monograph on
the Oligocene mammals of western Gansu, B. Bohlin (1946) divided what he regarded
as the middle-upper Oligocene into four levels. They are the following (in ascending
order): Hsanda-Gol, Wu-tao-ya-yu, Shih-chiang-tzi-ku, and Yindirte (Taben-buluk).
The distribution of the fossil mammals in these levels is as follows. Tataromys is
known from Hsanda-Gol to Shih-chiang-tzi-ku; Leptotataromys is restricted to Shih-
chiang-tzi-ku level in Gansu. Recently, several species, including the type species of
Tataromys, were found at Ulantatar locality, Nei Mongol, which was believed to be
comparable to Shih-chiang-tzi-ku (Huang X.s., 1987). Tataromys suni was first estab-
lished at Xiejia locality of lower Miocene (vide infra), but later was recorded in
Ikebulage Formation of late Oligocene, Nei Mongol (Wang B.y., pers. com.). The new
form, Metexallerix, is a very advanced form of brachyericines. It shares so many
532
Table z. The Neogene mammalian succession of China and its correlation with that of Europe.
! u R 0 p E c H I A ( continent.al)
Rogl and Steininger, 1983 Fablb.lsch,
•
Moin, Li c. k. et aJ..,
1976 l'fn Q 1 u z. x. ( t h i s paper)
1984
M a r i n • Continental . Represe.nta- Referred assemblages Moat characteristic roms
~:I! 0
"' tive Loce..l
Epoch• Stage• Rio- Faunas
Stages 0 Stages
zones
.
j~ MN) 0,
]]
".
Pleiat. Calabrian 'S
2 Villa£ran- ihewanian Shagou (38)
~ !i! VI I ~ IUYi!!!mit.SI~Ih !ll.!!!£1W.~
chian 17
E Piaoenzian 16 ouhean Youhe (36) Dongyaozi tou (37) I ~ youheensis. Himomxs~
I ------ 15 Jingle (1) (35)
. Zenel.ean Ruscinian
i Jinglean v Ge.ozhuang (34)
..j Ii :Hic:!;:er!!],!teg, Paracamel.us
5 i 14 ~
Messinian 13 Ert..te (33) '~minor. Loll:hocr!cetul!
. Turolian 12 Baodean Be.ode (31) Lufeng (32) etc, ~ Sinetherium.
... 11
Tortonian IV
---.-- Hezheng (26), Biru (27),
Bahe (25)
10
. Valle8ian
10
. Bahean
Wuzhong (28), Zho;;rng (29)
Wangdaifuliang (30 I
Di!!Q!ZQCU!d!, 1 Ninma therium
.-<
9 Qaidem (23) Amuwusu (24) ~ tsaidamensis
I ~
. ----- ---
8 " Tunggur ( 18) Platybelodon ~ Oioceros m!£1
. m>:"t~:~;:mf~- !IHj
'1\mggurian 1~7'jehuigou Xiaolongtan (22) enepqtberi umll, Pal.aeotragus
" .-< Serra118l.l- Astaracian 7 III
ian .
. "' Tongxin ( 13) '
15 6 mi;a~l~~~u~i6rugou Platvbelodon tongxinensis, Kubangchoerus 1
"'
-.< ------ --- 0 -------- : - -
0 Langhian
• 5 Shan wang ( 11) Xisodian ( 12) ne&aceratheriupl. Chalicotheriurp. Bl:!Yl!i!
0
II
0 Demoerig!:!i,Qdon, j&rcatheriy
&rdigelian Orleanian 4 Shanvangian Sihc>ng (9) Pangshan (10)
-.<
-.<
20
.. Zhangjisping 9,Qmabg:Y!eD,um, ~
~~~,;;) (t~~ Hsishui (Ts-
(6)
.-< 3
:E :E I
~
- --- -----· --- '-------

2 XieJis(J) Anlihsi ( 4) , Wa.£e.ngyongzi !Jericetodon. ~ 1!!.Q!.
(5
.. ~uitanian Agenian Xiejian
lAnzhou (1) SUosuoquan (2) Metepllerix. ~(primitive f'orms)
1
0'1
(..) L......-
(..)
(11
w
...
llt.l!l:l:r.•l r.,.. r{f ~l] illlf
Fig. 1. Important Neogene mammal localities of China (numbered as in table 2.).

common features with Exallerix of the Hsanda-Gol fauna that there is no doubt that it
was derived from the latter. However, the difference between these two forms seems
much larger than among the other lineages evolving from the middle to late
Oligocene. This had led us to think that this form, and hence the Lanzhou local fauna
as a whole, may be of Miocene age. In favor of this view is the fact that the known
distribution of this subfamily on other continents is restricted to the Miocene. The
American form comparable in evolutionary level to Metexallerix is Brachyerix, which
first appeared in North America in the latter half of the Arikareean age, at about
2.1 Ma. This is also the earliest record of that subfamily in North America. In Europe
there is only one distantly related form, Dimylechinus, which is known from MN
zone 1 (Saulcet, France). Judging by its tooth formula and morphology, Dimylechinus
cannot be attributed to the same lineage as the other members of the Brachyericinae;
but it is the only representative of that subfamily in Europe. Dimylechinus is much
more primitive than the Lanzhou species. All this has inclined us to think that the
local fauna of Lanzhou might be of the earliest Miocene age.
In the northern part of the Dzungar Basin, Xinjiang, from the Suosuoquan
Formation, which lies unconformably between the Wulungu Formation (Cretaceous)
and Halamagai Formation (middle Miocene), a local fauna was found. According to
Tong Y.s. (1987), the fauna includes Sinolagomys and Tachyoryctoides. Sinolagomys is
very common there and seems morphologically more advanced than similar specimens
found in the Taben-buluk area. The inner folds of their upper molars are very deep,
and the talonids of the lower molars are very wide, not narrower than the trigonids. In
addition, Metexallerix, very similar to that of Lanzhou, is also present (Mckenna, M.,
pers. com., 1988). This local fauna is very similar to that of Lanzhou.
Xiejia Fauna
In 1978 a local fauna of predominantly small mammals was discovered in the

Xiejia Formation of the Xining Basin of Qinghai Province. Li C.k. and Qiu Z.d. identi-
fied the following forms in 1980:
Sinolagomys pachygnathus
Sciurid sp.
Eucricetodon youngi
Plesiosminthus xiningensis
Plesiosminthus huangshuiensis
Plesiosminthus lajeensis
Tataromys suni
Tataromys sp.
Tachyoryctoides kokonorensis
Mustelidae gen. et sp. indet.
Brachypotherium sp.
Oioceros(?) xiejianensis
Most of these forms are highly specialized over their counterparts of the middle
and late Oligocene. There are few forms common to both Asia and Europe in this
fauna. Plesiosminthus was previously considered a widespread genus occurring in the
Holarctic region. However, recently Wang B.y. (1985) has pointed out that the
Chinese "Plesiosminthus" should belong to a separate lineage, phylogenetically distinct
from those of Europe and North America. She revived its original name,
Parasminthus, accordingly. The only genera common to both continents are, there-
fore, Eucricetodon and Brachypotherium. As far as the evolutionary level of
Eucricetodon youngi is concerned, Li and Qiu first (1980) considered it comparable to
the European Eucricetodon infralactorensis, but later (1984) they compared it with
Eucricetodon aquitanicus. The European FAD of E. infralactorensis is in MN zone 3,
while the FAD of E. aguitanicus is restricted to. MN zone Z (Ringeade, 1979). In fact,
these two European species are very similar in morphology. According to L. de Bonis
(1973), they differ mainly in size. Eucricetodon youngi is small for its advanced
morphology. Its Ml/ has a length of 1.77 mm; that for E. aquitanicus is 2..32. mm
(mean); and for E. infralactorensis is Z.SZ mm (mean), according to de Bonis' data.
636
The small size, in combination with some other peculiarities, like transverse direction
of the intersinus on the upper molars, led us to think that Eucricetodon youngi should
belong to a lineage different from the European E. collatus-E. infralactorensis
lineage. In spite of this,~· youngi could serve as an indicator of early Miocene age for
the deposits bearing it. Brachypotherium is represented only by a Mt IV and some
fragmentary teeth. Li and Qiu based their identification primarily on the Mt IV.
Reexamination of the material has revealed that the teeth are definitely different
from the teeth of typical Brachypotherium. The broken P4/ lacks any of the essential
characters of that genus, such as the antecrochet, the separation of the two trans-
verse lophs, and the development of the lingual cingulum. On the contrary, it is rather
high-crowned, especially as seen from the lingual side. The medisinus is blocked
lingually at the mid-height of the crown. The posterior cingulum is particularly devel-
oped, forming a conspicuous postfossette. The lower premolars are molariform, but
very small in size. These features are more characteristic for primitive elasmotheres
than for brachypotheres. The Mt IV is characterized by comparative robustness and
large lateral and plantar tuberosities on its proximal end. None of the known
brachypothere Mt IV conform with that from Xiejia, especially as judged by their
facets articulating with cuboid and Mt ill. In size, the Xiejia Mt IV is close to those of
Diaceratherium lemenense-aginense of Paulhiac and Laugnac (de Bonis, 1973, p. 137).
As regards "Oioceros" of the Xiejia Formation, according to Chen G.f. (1988) who has
reviewed the Chinese Oioceros, it may belong to a new genus, quite different from all
the other known Oioceros material.
The absence of proboscidean fossils and the typical Oligocene forms, like
Tataromys, Leptotataromys, Desmatolagus, etc., seems to indicate that the local
fauna from the Xiejia Formation is younger than the above described Lanzhou local
fauna, but older than the FAD of Asian proboscideans.
Fossils of similar age were also found in the southwest corner of Dzungar Basin
(Anjihai) of Xinjian Uygur Autonomous Region. From the middle part of the Shawan
Formation (the so-called brown beds), thick continental deposits, widely distributed
along the northern slope of the Tianshan Mountains, Dzungariotherium and
"Lophiomeryx" were reported (Chiu C.s., 1965, 1973). The first taxon is represented
by a huge skull in association with its lower jaw, probably the largest giant rhinoceros
ever found. It is also very advanced in morphology. The "Lophiomeryx" specimen is so
high-crowned and progressive that its identification should be considered tentative. A
new genus is likely to be established for it.
As early as 1962 Li C.k. described some fossils from a layer of claystones inter-
calated in the so-called Hannoba (Andersson's Han Jo Pa) basalt. The locality is called
Wafangyingzi, for a small village in northwest Zhangbei County, Hebei Province. The
fossils were identified as Monosaulax changpeiensis, a new species of that genus, and
Titanomys sp. The former is represented by a well preserved lower jaw, while the
latter is only a P4/. Since Monosaulax is mainly found in the Barstovian in North
America, the geological age of the Zhangbei locality was tentatively assigned to the
Miocene. Reexamination of the fossils has revealed the following. First, it is more
rational to associate the lower jaw of the beaver with the European genus Steneofiber
than with the American genus Monosaulax. Li has noticed some differences between
the American Monosaulax and that of the Zhangbei material; for example, the
comparatively low crown in the latter. We may add more: additional anteromost
fossettes on the lower molars are only occasionally present in Monosaulax; further-
more, the labial striids of the lower molars are evidently shorter than in the American
Monosaula~ (compare with fig. 73, Stirton, 1935); and the protoconids and hypoconids
are more rounded than in the typical Monosaulax. Steneofiber is known from late
Oligocene (Cournon) to at least middle Miocene (MN zone 5, Pontlevoy) in Europe.
According to Ginsburg {1971), it comprises a series of species increasing in size from
the Stampian to the Burdigalian Stage. Judging by its size, the Zhangbei lower jaw
stands just between those of Cournon and Saint-Gerand-le-Puy. Furthermore, in early
wear the additional anteromost fossettes are always present in Steneofiber. It is
difficult to ascertain the true affinity of the P4/ of the "Titanomys" sp. for the
536
moment, because the material is too poor. However, this P4/ has labial roots very
much reduced plus rather high crown and deep lingual groove. All this shows that it is
really comparable with the European Titanomys. Titanomys is a typical Aquitanian
ochotonid of Europe. The Zhangbei specimen conforms best with the early form, .:£..
visenoviensis. In conclusion, the two forms from Zhangbei suggest one and the same
geological age: early Miocene.
Zbanjiaping Fauna
In 1987 a very interesting fauna was found in a locality called Zhanjiaping, some
30 km north of the capital of Gansu Province, Lanzhou. The fossil-bearing deposits
are yellow coarse sandstones. Stratigraphically, they constitute the basal part of the
third formation of a thick continental series of the Lanzhou Basin. The formation is
called by the local geologists Xianshuihe Formation, a name first created by Young
C.c. and Bien M.n. in 1936. The material is being studied, but the preliminary identi-
fication reveals the following forms:
Tachyoryctoides sp., a left lower jaw without teeth. It is probably the largest speci-
men ever found for this genus. The breadth and length of the cross section of the
incisor are 5.2 mm and 6.1 mm (measurements for.:!:.· pachyrfcathus: 4.9 and 5.7; .:!:.·
kokonorensis: 4.3 and 4.7). The length of M/1-M/3 is 16 mm Tor.:!:.· pachygnathus it is
13-14 mm).
Hyaenodon sp., fragments of several premolars and a pair of upper carnassials (M2/) of
huge size (length more than 50 mm).
Proboscidea gen. et sp. indet., some fragments of tusks.
?Phyllotillon sp., an edentuous skull and a right M/3. The size of the M/3 is
44 by 20.7 mm, thus comparable to Phyllotillon naricus or Borissiakia betpakdalensis.
Aprotodon sp., an enormously widened symphysis with tusks (I/2) and several isolated
tusks.
Indricotheriidae gen. et sp. indet., a left scaphoid and fragments of teeth.
It is very interesting to note that, except for Tachyoryctoides and Hyaenodon,

the other forms have also been reported from the Bugti beds. The presence of the
proboscidean fossils and the absence of advanced forms of later age {like the progres-
sive cricetids, etc.) tend to show that this fauna may be correlated with MN zone 3 of
the European scale. In the zonation proposed by P. Mein, the FAD of proboscideans
and progressive cricetids were placed in MN zone 4a, but T. Antunes (1984) argued
that the proboscideans appeared in Portugal in MN zone 3b, while the progressive
cricetids (Democricetodon, Megacricetodon, etc.) made their first appearance in MN
zone 4. The data presented from China so far have tended to conform with those from
Portugal.
In 1986 we found another fauna of about the same age from the Linxia Basin at a
locality called Jiaozigou. This site is in a ravine some 10 km west of the Dongxiang
Autonomous County, about 50 km southwest of Lanzhou. The fauna was discovered in
the second member of the Linxia Formation, which had previously been considered
late Miocene in age. According to our preliminary study (Qiu Z.x. et al., in prepara-
tion), the fauna consists of Dzungariotherium orgosense, represented by several lower
premolars and molars and fragments of upper teeth; Caenopinae gen. et sp. indet.,
represented by only a lower molar; Paraentelodon sp. nov., represented by upper and
lower jaws with canines and cheek teeth, and a fragment of a tusk of Gomphotherium
sp. The association of the latter with the other specimens needs to be verified, but
the preservation of the proboscidean tusk is about the same as that of one of the upper
jaws of Paraentelodon. Again, this fauna is characterized by the combination of the
proboscidean fossils with highly specialized survivors of the Oligocene.
537
The fossils collected by B. Bohlin from the uppermost part of the thick and
tectonically disturbed deposits of Taben-buluk in 1931 and 1932 could also be included
in this faunal level. They were found in a number of small localities, the most
important of which were localities near Hsi-shui, Tien-chiang-tzi-ku, and Yindirte.
The forms commonly found there were: aff. Gomphotherium connexus, Sayimys
obliquidens, Kansupithecus (without species name), and fragmentary material of
Cervidae, Bovidae, and Rhinocerotidae. The Gomphotherium material is very
primitive in morphology. Its M3/ is very small, with only three lophs and a talon,
smaller and more primitive than the type specimen of the species from the Xining
Basin and those of the Bugti fauna of Pakistan.
Sihong Fauna
Systematic investigation and excavation of the Sihong fauna started in 1981.

The fossiliferous area is situated some 150 km northwest of Nanjing, the capital of
Jiangsu Province. Fossils were found in a number of sites. The fossil-bearing layer
consists of conglomeratic sands belonging to the Xiacaowan Formation, which is alto-
gether 90 m thick. Except for a few skull and jaw materials, the fossils are predomi-
nantly isolated teeth and bone fragments. In 1983 a preliminary list of fossils found in
that area was given by Li C.k. et al.; 65 taxa were recorded, among which 47 belonged
to mammals. A thorough subsequent study has shown that many of the first identifi-
cation must be reconsidered or corrected. The specimens so far restudied are:
Aplodontidae (Qiu Z.d., 1987)

Ansomyinae Qiu, 1987
Ansomys orientalis Qiu, 1987
Sciuridae (Qiu Z.d. and Lin Y.p., 1986)
Petauristinae
Parapetaurista tenurugosa Qiu et Lin, 1986
Shuanggouia lui Qiu et Lin, 1986
Sciurinae
Eutamias sihongensis Qiu et Lin, 1986
Plesiosciurus sinensis Qiu et Lin, 1986
Castoridae (Zhou M.z. and Li C.k., 1978)
Youngofiber sinensis (Young, 1955)
Gliridae (Wu W.y., 1986)
Microdyromys orientalis Wu, 1986
Primates
Dionysophithecus shuangouensis Li, 197 8
Platodontopithecus jianghuaiensis Gu et Lin, 1983
Carnivora (Qiu Z.x. and Gu Y.m., 1986)
Semigenetta huaiheensis Qiu et Gu, 1986
Pseudaelurus c£. lorteti
The presence of the following genera in the Sihong fauna is more or less certain,
though a detailed study of them is still not finished: Sayimys, Diatomys, Mega-
cricetodon, Democricetodon, Spanocricetodon, Anchitherium, Dorcatherium,
Lagomeryx, and Stephanocemas. The affinity of the proboscideans is still not clear,
but they are definitely present in this fauna. Two originally identified genera,
Protictitherium and Palaeotragus, must be discarded. Examination of the specimens
assigned to them has shown that the first may be an upper jaw of Semigenetta, while
the latter might belong to Palaeomeryx.
Taken as a whole, the fauna seems to be a little later in age than Wintershof-
West. Qiu Z.d. pointed out that the aplodontids of Sihong are morphologically more
advanced than those of Wintershof-West. While studying the carnivores of that fauna,
Qiu Z.x. and Gu Y.m. noted also that Semigenetta of Sihong is a little more advanced
than that of Wintershof-West. The abundance of Dorcatherium material here tends
also to support this point of view, because Dorcatherium appeared in Europe from MN
zone 4. It is rather unexpected to find a beaver so large and advanced as Youngofiber
in the Sihong fauna. The early and middle Miocene beavers in Europe were all smaller
and more primitive than Youngofiber.
538
From Fangshan of Jiangning County, some 20 km south of Nanjing, the horse
Anchitherium cf. aurelienense (Chow M.c. and Hu C.k., 1956), the rodent Spano-
cricetodon ningensis (Li C.k., 1977), and some ochotonid and cervid fossils were
found. The fossil-bearing layer is unquestionably overlain by a basalt dated about
14 Ma (Shao J .j. et al., 1987). This fauna may belong to the same faunal zone as
Sihong.
Shanwang Fauua
Shanwang is one of the few Chinese Miocene fossil localities discovered before
the founding of the People's Republic. It has been famous for its exquisite specimens
of various groups of animals and plants preserved in diatomaceous deposits. The
locality lies about ZO km east of Linqu County seat, in the middle part of Shandong
Province. Yan D.f. et al. (1983) reviewed current knowledge of the site. They listed
Z1 mammalian forms. In recent years some new taxa have been added to the faunal
list and restudy of some previously known forms were also made. Unfortunately, its
radiometric age is still not convincingly settled, although there is plenty of basalt in
association with the fossiliferous layers. As far as the mammals are concerned, the
following are securely identified:
Chiroptera
Shanwangia unexpectula Young, 1977
Rodentia
Ansomys shanwangensis Qiu et Sun, 1988 (aplodontid)
Plesiosciurus aff. sinensis (sciurid)
Meinia asiatica Qiu, 1981 (petauristid)
'i5'fat0mys shantungensis Li, 1974 (geomyid?)
Carnivora
Hemicyon (Phoberocyon) youngi (Chen, 1981)
Amphicyon confucianus Young, 1937
Ursavus orientalis Qiu et al., 1985
Thaumastocyoninae gen. et sp. indet.
Proboscidea gen. et sp. indet.
Perissodactyla
"Palaeotapirus" xiejiaheensis Xie, 1979
Plesiaceratherium gracile Young, 1937
Brachypotherium sp. (an unstudied skeleton)
Chalicotherium grande (Mtm and phalanges, unstudied)
Artiodactyla
Palaeomeryx tricornis Qiu et al., 1985
Lagomeryx spp. (to be studied)
Suinae (to be studied)
Anchitherium seems to be absent from this fauna. The referral of a distal end of
metapodial to this genus by Yan et al. (1983) was in error as it proves to belong to
"Palaeotapirus".
L. Ginsburg and E. Heinz (1960) believed Palaeomeryx increased in size regularly

from its first appearance in MN zone 3 to zone 9. Qiu et al. (1985) pointed out that
the size of the teeth of the Shanwang Palaeomeryx was comparable to that of MN
zone 5 (Pont Levoy). While studying the aplodontids, Qiu Z.d. and Sun B. came to the
same conclusion as regards the age assignment of the Shanwang fauna. Ansomys
shanwangensis is evidently more advanced in tooth morphology than Ansomys
orientalis of Sihong, according to Qiu and Sun (1988). The new record of foot bones of
Chalicotherium grande strongly supports this age assignment, because the species in
question appeared in Europe also in MN zone 5.
Hemicyon (Phoberocyon) youngi was also found in a locality near Xiaodian

village of Zhongxiang County, Hubei Province. In association with this carnivore there
were also an upper molar of "Macaca" youngi Gu, 1980, and some teeth of Cervidae.
Based on the common presence of H. youngi, this site may be of the same age as
539
Shanwang. The occurrence of a Macaca in a locality of Hemicyon seems highly per-
plexing and has caused much dispute. M. Pickford believes that this tooth should
belong to a tayassuid genus (pers. comm., 1988).
Tongxin Fauna
During the 1970s, news about "dragon-bones" from the Tongxin area of Ningxia
Hui Autonomous Region often came to us. As a result of cooperative effort between
the Institute of Vertebrate Paleontology and Paleoanthropology, the Beijing Museum
of Natural History, and the Bureau of Geology of the region, a collection of
mammalian fossils was made. Some specimens have been published, but the majority
of the material is still under study. Specimens described so far include a single tooth
of Pliopithecus sp. (Qiu Z.x. and Guan J., 1986), a sample of teeth (about 50) of
Platybelodon tongxinensis (Ye J. and Jia H., 1986), a skull and several incomplete
lower jaws of Kubanochoerus lantienensis (Qiu Z.x. et al., 1988), and skull and lower
jaws of Percrocuta primordialis (Qiu Z.x. et al., 1988). Undescribed material includes
a skull of Sansanosmilus, a partial skull and teeth of Gobicyon, jaw of a schizothere
(Ancylotherium or Metaschizotherium), skulls and jaws of at least two types of rhinoc-
eros (an elasmothere and an acerathere), and horn-cores of Eotragus. Among small
mammals there are many isolated teeth of ochotonids.
Taken as a whole, the Tongxin fauna is very close to that of Belometchetskaya

of the Soviet Union, which is now commonly regarded comparable to MN zone 6 (or
even 5) of the European zonation.
The type specimen of Kubanochoerus lantienensis was found in the Koujiacun

Formation near the village of Koujiacun, some 10 km east of Xi-an, the capital of
Shaanxi Province. Two other forms of listriodonts were also reported from this site:
Kubanochoerus gigas and "Listriodon" intermedius (Liu T.s. and Li Y.q., 1963). The
locality was referred originally to middle Miocene, that is, earlier than the age of
Tung-gur, but later (in 1978) it was regarded as directly comparable with Tung-gur in
age (Zhou M.z., 1978). Based on the new listriodont materials in Tongxin and Tung-gur
areas, we now prefer to correlate the Koujiacun locality with Tongxin rather than with
Tung-gur. While Tongxin has the same species of Kubanochoerus as Koujiacun,
Tung-gur has probably a more advanced and larger species, although the material from
Tung-gur is very poor (only an incisor). For the same reason, we would assign the type
locality of Kubanochoerus gigas at Quantougou near the village Xiejia in Yongdong
County, Gansu, to the Tongxin faunal level.
It is a very difficult problem to assign an age to the Jiulongkou fauna from

Cixian, Hebei. This locality was first mentioned in Young's paper on the Shanwang
fauna. Hu C.k. (1959) described a lower jaw of "Macrotherium" cf. brevirostris.
Later, Chen G.f. and Wu W.y. (1976) gave a full report on this fauna, describing
altogether 16 forms. Among them the most important are:
Carnivora
Sansanosmilus palmidens
Percrocuta hobeiensis
Perissodactyla
"Macrotherium" sp.
Dicerorhinus cixianensis
Plesiaceratherium gracile
Chilotherium sp.
Artiodactyla
Palaeomeryx sp.
Oioceros(?) jiulongkouensis
0. (?) robustus
Q. (?) stenocephalus
Reexamination has revealed the following: Percrocuta hobeiensis is at most a

subspecies of R_. miocenica, as pointed out by C. Howell and G. Petter in 1985. Chen
540
and Wu's identification of Sansanosmilus is confirmed. The specimens of Jiulongkou
conform very well with those described by L. Ginsburg in 1961. The lower jaw of
"Macrotherium" cf. brevirostris described by Hu C.k. is apparently more progressive
than those of Chalicotherium grande, which was so amply demonstrated by H. Zapfe in
1979. Zapfe's material came from Neudorf-Spalte an der March, Czechslovakia, now
assigned to MN 5 (Mein, 1981). The progressiveness of the Jiulongkou lower jaw is
clearly shown in its much enlarged angular lobe. Dicerorhinus cixianensis is at about
the same evolutionary level as D. sansaniensis (Lartetotherium, after L. Ginsburg,
1974), as shown by their conformity in both morphology and size. The chilothere
specimens of Jiulongkou are really similar to Chilotherium of the HipParion fauna.
They differ from the latter by less reduction of the DP1/, generally smaller size,
comparatively shallow nasal-notch, broader nasal bones with lateral borders turning
down, and lesser shifting of the preorbital foramina onto the dorsal surface of the
maxillary bones. In general, all the above distinctions indicate that the Jiulongkou
chilothere species is more primitive than those associated with Hipparion. In fact,
this is the oldest occurrence of that genus in China, and probably in the world. The
other material of rhinoceroses identified by Chen and Wu was based mainly on unchar-
acteristic specimens, and, therefore, cannot be confirmed. The Palaeomeryx speci-
mens are hardly separable from those described from Shanwang (Qiu z.x. et al.,
1985). The presence of rich material of "Oioceros" is noteworthy. Chen and Wu
established three new species. It is remarkable that in only one skull of "0."
jiulongkouensis a pair of knob-like hom-cores has been observed.
Whether the Jiulongkou fauna is contemporary with Tongxin is difficult to

decide. They share only a few common taxa: Sansanosmilus and probably Percrocuta
miocenica. Taxa that are characteristic for the Tongxin fauna are strangely absent in
the Jiulongkou fauna. For example, Kubanochoerus, Platybelodon, elasmotheres, and
aceratheres occur at Tongxin, while in Jiulongkou there are other forms, like
chalicotheres, dicerorhine, chilothere, Palaeomeryx, and many "Oioceros". However,
as far as their age is concerned, both seem to belong to zone MN 6, if we compare
them separately with the European faunas. If we attach greater importance to new
forms rather than retention of old ones, we would be inclined to consider Jiulongkou a
little younger than Tongxin. The difference is hardly to be explained by ecological
factors, since both faunas have high-crowned ungulates.
Qiu Z.d. et al. (1981) described some fossil mammals, mainly micromammals,
from the Xining Basin, Qinghai Province. The fossil-bearing deposits ranged from the
base of the Xiejia Formation upward to the Chetougou and Xianshuihe Formation.
From the Chetougou Formation a new species of Megacricetodon was described: M.
sinensis. In addition, Protalactaga tungurensis was also found. Plesiodipus leei and a
new species of Alloptox, plus some poorly defined large mammals, such as
Gomphotherium connexus, G. wiman, Micromeryx, "Oioceros", and Stephanocemas and
a new species of Kubanochoerus, K. minheensis were reported from the Xianshuihe
Formation. Judging by the small size of the Kubanochoerus and the absence of char-
acteristic elements of the Tung-gur fauna, like Platybelodon, Palaeotragus, etc., it
seems plausible to restrict the age of the fossiliferous deposits of the Xining Basin to
the interval spanning from the Xiejia to the Tongxin fauna. At present no closer
comparison can be made because of the fragmentary nature of the large mammals
collected.
Lengsbuigou Fauna
This locality is situated about ZO km northeast of Xi-an, Shaanxi Province. A

small fauna was found in the Lengshuigou Formation and described by Zhai R.j., Chang
H.s., and Zhai R.j. (1978). The fauna consists of:
Proboscidea
Selenolophodon spectabilis Zhang et Zhai, 197 8
Perissodactyla
Hispanotherium lintungense Zhai, 1978
541
Artiodactyla
Listriodon lishanensis Li et Wu, 1978
Listriodon sp.
Palaeomeryx sp.
Palaeotragus sp.
"Oioceros" lishanensis Li et Wu, 1978
Ye J. and Jia H. (1986) held the opinion that the new genus Selenolophodon of
Lengshuigou was nothing but a species of Platybelodon intermediate in morphology
between P. tongxinensis and f.• grangeri. The other forms all seem to indicate that
the Lengshuigou fauna should be a little younger than Tongxin. Instead of
Kubanochoerus, the true lophodont genus Listriodon appeared here. The Oioceros skull
bears true horn-cores typical of that genus, though small in size. We notice also the
first appearance of the true giraffe, Palaeotragus. There is only one form,
Palaeomeryx, which is little changed and is very similar to the specimens discovered
from Cixian and even Shanwang. The Hispanotherium of Lengshuigou is larger in size
and more progressive in morphology than the type specimen of the genus (in Spain,
zone MN 5, see Antunes and Ginsburg, 1983) and the specimens we found from
Tongxin. Its large size can only be judged by the breadth of the teeth, since the skull
is of a very old individual. The molars of the Lengshuigou form have a very strong
constriction of the protocones, enlarged antecrochets, especially on M3/, and marked
enamel crenulation.
Paradoxically, the Lengshuigou fauna has long been considered earlier than the
Koujiacun fauna in age (Zhou M.z., 1978). It appears that this happened partly through
an erroneous association of fossils of different ages and partly because of inexact
correlation of the deposits of different areas. However, from the paleontological
viewpoint, it is quite clear that the Lengshuigou fauna lies just between those of
Tongxin and Tung-gur. We have little doubt about it, after having compared the
materials of the three faunas.
Tuug-gur Fauna
Tung-gur is the only locality in China that has been well known to the world
since its sensational discovery in 19Z8. A recent campaign of renewed excavation and
restudy has given very encouraging results (Qiu z.x. et al., 1988). The number of the
fossil mammals was increased from Z9 to 59. The majority of the new forms are
micromammals that are presently being studied. The following is a preliminary list of
the fossil mammals so far known from Tung-gur:
Insectivora
Erinaceinae gen. et sp. indet. (1-3)
Desmanella sp.
Talpidae gen. et sp. indet.
Soricidae gen. et sp. indet.
Chiroptera gen. et sp. indet.
Rodentia
Ansomyinae gen. et sp. nov.
cf. Atlantoxerus sp.
Sciurus sp.
Spermophilinus sp.
Anchiterhiomys tungurensis (Stirton, 1934)
"Monosaulax" tungurensis Li, 1963
?Microdyromys sp.
Keramidomys sp.
Leptodontomys sp.
"Protalactaga" tungurensis Wood, 1936
Protalactaga cf. grabaui Young, 19Z7
Paralactaga sp.
Plesiodipus leei Young, 19Z7
Cricetodon sp.
542
Megacricetodon sp.
Democricetodon sp.
cf. Cotimus sp.
Lagomorpha
Alloptox gobiensis (Young, 1931)
Bellatona forsythamajori Dawson, 1961
Ochotonidae gen. et sp. indet.
Carnivora
Gobicyon macrognathus Colbert, 1939
Pseudarctos sp.
Hemicyon teilhardi Colbert, 1939
Amphicyon tairumensis Colbert, 1939
Leptarctos neimonguensis Zhai, 1963
Melodon sp.
Mionictis sp.
Martes sp.
Tungurictis spocki Colbert, 1939
Percrocuta tungurensis (Colbert, 1939)
Metailurus mongoliensis Colbert, 1939
Machairodus sp.
Proboscidea
Platybelodon grangeri Osborn, 1929
Serridentinus gobiensis Osborn et Granger, 1932
Zygolophodon sp.
Perissodactyla
Anchitherium gobiensis Colbert, 1939
Chalicotherium brevirostris (Colbert, 1939)
Rhinocerotidae gen. et sp. indet. (1-3)
Artiodactyla
Listriodon mongoliensis Colbert, 1934
Kubanochoerus sp.
Stephanocemas thomsoni Colbert, 1936
Dicrocerus sp.
Dicrocerus grangeri Colbert, 1936
Micromeryx sp.
Lagomeryx triacuminatus (Colbert, 1936)
Euprox sp.
Palaeotragus tungurensis Colbert, 1936
"Oioceros" grangeri Pilgrim, 1934*
"Oioceros" noverca Pilgrim, 1934
In Xinjiang Uygur Autonomous Region a fauna of similar age was found recently
from the northern part of the Dzungar Basin. The localities are situated along the
northeast bank of the Urungu River. The fauna was preliminarily published by Tong
Y.s. (1987). The fossil-bearing deposits are mainly yellow sandstones intercalated with
gray-green clays. The whole series of the intercalated layers is called Halamagai
Formation. In addition to the forms in common with Tung-gur, such as Platybelodon,
Anchitherium, Stephanocemas, some odd forms have also been found: a new genus
representing the first record of the Mylagaulidae in Eurasia and a new species of
Atlantoxerus.
A small faunal assemblage reported by Yan D.f. (1979) from a locality called
Erlanggang, Fangxian, Hubei, may to be attributed evidently to the same faunal level
as Tung-gur. The assemblage consists of Zygolophodon neimonguensis, Anchitherium
aurelianense, Tesselodon fangxianensis (an elasmothere), Listridon robustus, and
"Oioceros" noverca.
*Based -
on "Oioceros" grangeri, M. Kohler (1987) erected a new genus Turcocerus,
while Chen Guanfang (1988) erected another one, Sinomioceros.
543
The locality Linyanshan, some ZO km north of Nanjing, Jiangsu, has yielded some
fragmentary teeth of Hyotherium cf. paleochoerus, Tetralophodon sp., and
Acerorhinus sp. (Bi Z.g. et al., 1977). These fossils were found in a layer about ZO m
lower than a basalt flow dated as 1Z.17 Ma by the K-Ar method (Shao J.j. et al., 1987).
The assignment of the Xiaolongtan fauna near Kaiyuan, Yunnan, is rather

controversial. Recently, Dong Wei (1987) reviewed the fauna and, according to him, it
consists of 11 forms, the main ones of which are Sivapithecus sp., Tapirus cf.
yunnanensis, Propotamochoerus parvulus, Dicoryphochoerus sp., and Tetralophodon
xiaolongtanensis. He placed the fauna in late Miocene (comparable with Vallesian),
but we are inclined to consider it belonging to the terminal part of the Anchitherium
fauna, that is, the latest middle Miocene.
Qaidam (Tsaid.am) Fauna
The Qaidam fauna was described by B. Bohlin in 1937. Unfortunately, Bohlin did
not report on the geology of the localities. Judging by the fact that Bohlin always
referred to Qaidam as a single fauna with Hipparion, the fossils he described in his
monograph were likely collected from a single stratigraphic unit. Now it is believed
that Bohlin's Qaidam fauna came from the upper Youshashan Formation (Li Y.t.,
1984), from which similar fossils were found during the 1950s and 1960s. This fauna is
characterized by association of survivors of the Anchitherium fauna and elements of
the Hipparion fauna. For example, genera of the first group are Stephanocemas,
Lagomeryx, and Dicrocerus; those of the second group are Hipparion, Tetralophodon,
and Ictitherium. In addition, there are also some primitive forms of the typical
Hipparion fauna, such as Acerorhinus tsaidamensis; there are also peculiar bovids, such
as Qurliqnoria, Olobulukia, Tsaidamotherium, etc.
In 1986, while prospecting the area surrounding Tung-gur, we found a small

locality with both Hipparion and Anchitherium teeth. The locality, called Amuwusu, is
situated 1Z km west of the railway station Zhurihe, which is about 150 km to the south
of the border city Erlian and about ZOO km southwest of Tung-gur. Typical Tungurian
forms were found there: Bellatona forsythmajori, Anchitheriomys, Protalactaga
tungurensis, and Dicrocerus. The presence of a species of the genus Prosiphneus in
this fauna is of particular interest.
Babe Fauna
Along the south bank of the Babe River in the Lantian area, beneath the cover of
a layer of red clay of Baode type (Lantian Formation) and above the Koujiacun
Formation, fluvial deposits (Babe Formation) consisting chiefly of yellow sands and
conglomerates are exposed. In the 1960s these rocks yielded a fauna slightly different
from the typical Baode Hipparion fauna. The revised fauna list, by the present author,
is given below:
Erinaceus sp.
Herpestinae gen. et sp. indet. (originally as Viverridae)
Miomachairodus sp. (orig. Crocuta eximia variabilis, a P4/)
Dinocrocuta macrodonta (Liu et al., 1978),
(orig. Crocuta macrodonta)
Tetralophodon exoletus Hopwood, 1935
Hipparion (Hippotherium) weihoense Liu et al., 1978
Hipparion (Hippotherium) chiai Liu et al., 1978
Acerorhinus sp. (orig. Chilotherium gracile)
Dicerorhinus ringstromi Hooijer, 1966 (orig. Dicerorhinus
orientalis + ?Brachypotherium sp.)
Chleuastochoerus stehlini (Schlosser, 1903)
Palaeotragus microdon (Koken, 1885)
Samotherium decipiens Bohlin, 1926 (orig.
Paleotragus decipiens)
544
Shaanxispira chowi Liu et al., 1978
Antelope sp.
Gazella sp.
It is noteworthy that the Hipparion species are of the primigenium type and the
carnivores are different from those found in the typical Baode Hipparion fauna.
Miomachairodus is a genus recorded from Eskihisar (middle Miocene) and Esme-
Aksakoy (Vallesian) of Turkey.
A Hipparion fauna with Dinocrocuta and Acerorhinus was also recently reported
from Hezheng County, Gansu (Qiu z.x. et al., 1987, 1988).
In the locality near Biru, Tibet, a similar fauna was found. Dinocrocuta, another
primitive species of Hipparion, H. xizangense, Brachyrhizomys hehoensis, and a small-
sized "Chilotherium" tanggulaense are recorded from Biru.
A locality in Wuzhong County, Ningxia, yielded Hipparion weihoense, along with

fossils of Tetralophodon cf. exoletus, Acerorhinus cf. tsaidamensis, and Qurliqnoria
cheni (Qiu z.x. et al., 1987). In correlative rocks at another locality in Zhongning
County, some 80 km southwest of the Wuzhong County, a large skull of the
elasmothere, Ningxiatherium, was reported in association with a Hipparion metapodial
(Chen G.f., 1977). Ningxiatherium is much more primitive than Sinotherium of the
Baode Hipparion fauna.
Recently, a locality near the village of Wangdaifuliang in Fugu County, Shaanxi,

yielded a large quantity of fossils. Wangdaifuliang lies on the westem bank of Huang
river, opposite to Baode, Shaanxi. Most of these fossils are chilothere skulls, no less
than ZOO, but there are also skulls of Hipparion, giraffids, gazellas, and cervids. The
most interesting specimens are jaws of Dinocrocuta and a lower jaw of Platybelodon.
The material is still being prepared, but the presence of the last two forms inclines us
to think that this fauna might well be older than that of Baode.
Baode Fauaa
This faunal assemblage is typical for the classical Hipparion fauna of the red
clay in North China. Such assemblages are widespread in north China, especially in its
central part, in the great loess plateau area. The fossils are so numerous and so
famous (as "dragon bones and teeth") that they attracted the attention of the first
paleontologists working in China and served as the main object of many monographs in
the first period of extensive paleontological research in China (19ZOs-1940s). A
thorough revision of the whole fauna in light of recent practices in mammal taxonomy
and biostratigraphy is badly needed and constitutes one of the main subjects of the
working plan of the author; but it is far beyond the scope of the present paper.
One recent advance in South China Hipparion fauna study should be mentioned
here. That is the discovery of the hominoid fossil-bearing locality at Lufeng in
Yunnan Province. The fossils discovered and accumulated from Lufeng since 1975 are
numerous in quantity and variety, representing probably no less than 50 species;
however, in most cases they are fragmentary. The bulk of the material is now under
study; the following mammals have been published from Lufeng:
Rodentia
Brachyrhizomys nagrii (Hinton, 1933)
B. cf. pilgrimi
B. tetracharax Flynn, 198Z
Lagomorpha
Alilepus longisionuosus Qiu et Han, 1986
Primates
Sinoadapis carnosus Wu et Pan, 1985 (incl.~·
shihuibaensis, Pan et Wu, 1986)
545
Lufengpithecus lufengensis (Xu et al., 1978)
(= Ramapithecus lufengensis Xu et al., 1978
+ Sivapithecus ywmanensis Xu et Lu, 1979)
Carnivora
Ursavus cf. depereti Schlosser, 190Z
Jndarctos sinensis Zdansky, 19Z4
Ursidae gen. et sp. indet.
Proputorius lufengensis Qi, 1983
Sivaonyx bathygnathus Lydekker, 1884
Lutra sp.
'ICtitilerium gaudryi Zdansky, 19 Z4
Machairodus fires (Qi, 1983) (orig. Epimachairodus fires)
Metailurus sp. (orig. Pseudaeilurus sp.)
Artiodactyla
Dorcabune progressus (Yan, 1978)
Yunnanotherium simplex Han, 1986
The fossils of Proboscidea and Perissodactyla have not been published yet,
although we know for certain that there are Hipparion, chilotheres, tapirs, and so on.
The majority of the Artiodactyla, especially Suidae and Bovidae, and the Rodentia
(Muridae, Cricetidae, ...) have not been published either. The Brachyrhizomys
material has so far provided the most precise age assignment. According to Flynn and
Qi (198Z), the age of the Lufeng fauna should be about 8 Ma.
Ertemte Fauna
The locality Ertemte, Nei Mongol Autonomous Region, was discovered by J.

Andersson in 1919, and was described and published by M. Schlosser (19Z4). Since most
of the material is micromammals, it is difficult to compare it directly with the classi-
cal Hipparion fauna of North China, which is known predominantly by large
mammals. In spite of this difficulty, the Ertemte fauna has always been considered a
little younger in age than the typical Hipparion fauna. A recent campaign to collect
more material utilizing the screen-washing method was carried out in 1980 by a joint
expedition of Chinese and West German paleontologists. Study of the procured
material from Ertemte is now under way, with several papers already published.
According to the preliminary results given by Fahlbusch et al. (1983), the fauna
consists of at least 51 mammalian taxa, many of which are new. Systematic study of
the single groups so far completed further substantiates the former assignment of its
age as a little younger than the typical Hipparion fauna (Storch and Qiu Z.d., 1983; Qiu
Z.d., 1985; Wu W.y., 1985; Fahlbusch, 1987; Storch, 1987; QiuZ.d., 1987).
Gaozhuang Fauna
Gaozhuang is a village of Yushe County, Shanxi Province, about 10 km southwest

of the county center. Around the village there are numerous sites that yielded a large
part of the fossil mammals of Teilhard de Chardin's "Zone ll" of the Yushe Group.
Since Teilhard de Chardin's "Zone II" has never been clearly defined and its faunal
contents are highly mixed with those from both his "Zone I" and "ill," we prefer to
designate this stratigraphic interval by the village name, Gaozhuang. There is no
doubt that the Gaozhuang fauna is radically different from the Baode fauna and even
the Ertemte fauna. It is marked by the earliest appearance of the canid, Nyctereutes,
and the camelid, Paracamelus, apparently from North America. Furthermore, there
are many new forms that are more advanced or specialized than their counterparts in
the Boade or Ertemte faunas; for example, Mammut borsoni, Stegodon zdanskyi,
Chasmaporthetes kani, Pliohyaena perierri orientalis, Agriotherium sp., Ursus sp., an
advanced form of Plesiogulo, Hipparion houfenense, Proboscidipparion pater, and many
varieties of Paracervulus. This fauna is now under intensive study by a joint party of
Chinese and American paleontologists. At present the Gaozhuang fauna is roughly
correlated with MN zones 14 and 15 in the European scale.
546
The local fauna discovered at a locality near Hefeng, Jingle County in northern
Shanxi, has always been directly correlated with Teilhard de Chardin's Yushe "Zone
n." The fauna consists of the type specimens of Hipparion houfenense, Gazella blacki,
and Antilospira licenti. A plate of a milk tooth of Elephas sp. and some cervid and
rhinoceros limb bones were also found there. In 1986 a new excavation yielded some
new forms. Among the large mammals Metailurus and Nyctereutes were found, while
micromammalian taxa Chardinomys sp., Prosiphneus sp., and Ungaromys sp. were
added. Except for Elephas, the other forms are all more or less comparable to those
of the Gaozhuang fauna. The provenance of the Elephas tooth is questioned. Until its
presence in this fauna is verified or disproved, the correlation of the Jingle fauna
cannot be properly settled. However, it is quite possible that the Jingle fauna may
represent a transitional faunal interval between the Gaozhuang and the younger Youhe
fauna.
Youhe Fauna
The Youhe fauna was described by Xue Xiangxu (1981). The most important
forms are Elephas youheensis, Hipparion houfenense, Sus subtriguetra, Cervavitus sp.,
Nyctereutes sinensis, Ochotonoides cf. complicidens, and Mimomys youhenicus. H.
houfenense was later transferred to Proboscidipparion pater by Qiu z.x. et al. (1987).
Along with the type specimen of M. youhenicus, there were also specimens of M.
orientalis and Mimomys sp., the latter of which was considered very close to M.
banchiaonicus in size and morphology. According to Zheng and Li (1986), the
Mimomys of Youhe is evidently correlated with the primitive forms of that genus in
Europe, like M. gracilis, M. occitanus (e.g., Csarnotian), M. stehlini, and M. polonicus
(e.g., lower Villafranchian). Elephas youheensis is really very primitive in character.
The other members of the fauna show primitive characters as well. The Youhe fauna
appears to be older than the Shagou or typical Nihewan assemblage and is thus corre-
lated with the early Villafranchian.
A similar fauna, the Dongyaozitou fauna, was discovered also in the lower part
of the late Cenozoic section beneath the Nihewan Formation in Yu County. Initially
the fauna was described by Tang Yingjun (1980). Among the fossils described by Tang
(1980) and Tang and Ji (1983) three forms definitely indicate an age earlier than the
typical Nihewan fauna. They are Zygolophodon sp. (probably Mammut borsoni),
Antilospira yuxianensis, a new species, but very similar to the type specimens of A.
licenti, and Palaeotragus progressus. From about the same level Cai Baoquan (1987)
obtained a very important micromammalian fauna including Mimomys orientalis,
Germanomys sp., Orientalomys sp., Prosiphneus sp., Ochotona cf. lagrelii, etc. The
fauna is still being studied, but it is clear that both the large and the small mammals
of this fauna represent a fauna earlier than Nihewan (s.s.). It is correlated with MN
zone 16, probably 16a, in the European mammal zonation.
Shagou (Nihewan s.a.) Fauna
The Nihewan fauna (s.s.) has been well known since it was described by Teilhard
de Chardin and Piveteau (1930). However, its geological age had not been accurately
defined until recently. Now it is generally considered roughly comparable with Senez-
Le Coupet (MN zone 18) faunas of Europe, but younger than St. Vallier (MN zone 17) in
age. As far as its position in relation to the Plio-Pleistocene boundary is concerned,
opinions differ widely. Li C.k. et al. (1984) placed the Q/N boundary just below the
Nihewan (s.s.). The absolute date of the boundary was thus believed to be around
1.8 Ma. Zheng S.h. and Li C.k. (1986) later placed the boundary at approximately
Z.4 Ma, below a so-called Dachai fauna characterized by Mimomys peii, which was
thought to be contemporaneous with MN zone 17 of Europe. Unfortunately, this
faunal zone is left uncharacterized by large mammals. The majority of specialists
who study loess and paleomagnetism hold the same opinion as Zheng and Li, based on
the first appearance of loess in China fixed at about Z.4 Ma. The current practice in
geochronology outside China tends to fix the Q/N boundary at the Olduvai event of
Matuyama Chron (1.87 to 1.67 Ma). This is apparently younger than the Le Coupet
fauna which is dated at 1.9Z Ma. So far there has been no authentic evidence demon-
547
strating that the Nihewan fauna (s.s.) is later than Le Coupet in age. Thus, the
Nihewan fauna (s.s.), at least its lower part, should belong in the Pliocene, if we
accept 1.8 or 1.6 Ma as the boundary between the Neogene and Quaternary.
PALEOZOOGEOGRAPIUC SETTING
It is well known that today Europe and the main part of Asia belong to one
zoogeographic region, the Palaearctic. However, how far can this be traced back to
the Neogene? Further, what kind of relationships existed between Asia and North
America during that period? And, eventually, how can we deal with south China,
which belongs to the Oriental Zoogeographic Region? So far, these problems have not
been properly addressed.
The recent boundary between the Palaearctic and the Oriental Regions goes
through the Huai River drainage area in the east and along the Himalayas in the west
within the territory of China. It seems that this boundary did not exist at the begin-
ning of the Neogene. The recent discovery of fossils attributed to the early Miocene
in the northwest part of China shows that at least central Asia was strongly isolated
zoogeographically. The majority of the Chinese early Miocene mammals were
endemic and restricted to central Asia. They are mainly the holdovers of Asian Oligo-
cene faunas, such as Sinolagomys, Tataromys, Tsaganomys, Tachyoryctoides. Though
ill-defined, the presence of some primitive bovids is beyond all question. Various
forms of the Indricotheriinae were the most representative element of the faunas
from middle Oligocene through early Miocene. Unlike the other endemic forms, they
spread westward to the Transylvanian area of Rumania, but never reached west-
central Europe. A grotesque entelodont of enormous size, Paraentelodon, has been
found both in Gansu (in prep.) and the Caucasus of the Soviet Union (Gahunia, 1964).
In west-central Europe only one genus, Entelodon, was found, which did not survive
middle Oligocene (its last record is in Heimersheim). Genera seemingly common to
both Europe and Asia include Eucricetodon, Plesiosminthus, Brachypotherium, and
probably a "Phyllotillon." However, none of these would necessarily be explained by
direct dispersal from Europe or vice versa. Primitive Oligocene Eucricetodon was
found in both Europe and Asia. Thus, the Asian forms could have evolved in place. As
to "Plesiosminthus," Wang B.y. (1985) has demonstrated that the Asian
"Plesiosminthus" was basically different from the European genus in the absence of
grooves on the upper incisors and the presence of four-rooted upper molars. In her
opinion, the Chinese "Plesiosminthus" should be referred to Parasminthus as supposed
by Bohlin (1946). The still undescribed schizothere could also have evolved from some
local species of schizotheres, which were very common in the Oligocene of central
Asia. Brachypotherium is unfortunately represented only by a Mt IV and tooth frag-
ments. Its identification cannot be taken for granted.
According to recent studies (Steininger et al., 1976), during the Egerian and
Eggenburgian ages the Paratethys was still actively connected with the Mediterranean
Sea. The Paratethys linked with the Mediterranean Sea through a seaway from
Budapest and Zagreb to the Adriatic. However, on the eastern side of the Paratethys
the Uralian Sea and Turgai Strait had retracted by the late Oligocene. Therefore, in
principle, a free exchange of animals by a route north of the Paratethys should have
been possible. But, in fact, no such thing occurred. The possible explanation would be
that some ecological factors impeded free exchange between the two regions. The
Oligocene fauna so far discovered from South China (Yunnan) differs only slightly
from that of North China. Giant rhinoceroses have been found in Yunnan as well.
Furthermore, the Bugti fauna of Pakistan, which is temporally only a little later than
the Chinese early Miocene faunas discussed above, shows some similarity to these
Chinese faunas. Therefore, the whole mainland of Asia, together with the southern
part of east Europe, might constitute one zoogeographic region prior to the
Gomphotherium datum. As for the relationships between Eurasia and North America,
what we would recognize is only some casual migration events between them. One of
the few examples is the possible emigration of the brachyericines from Asia to North
America. As a result of the foregoing analysis, we have come to the conclusion that
548
during the early Miocene, prior to the Gomphotherium datum, Eurasia could be sub-
divided into two zoogeographic regions of first order: the west-central European
region and the Dinarid-Asian Giant Rhinoceros region.
With the almost contemporaneous immigration of Anchitherium from North

America and Gomphotherium from Africa the general paleozoogeographic pattern of
Eurasia changed. Probably the barrier which had impeded the giant rhinoceros to
traverse along the southern bank of the Paratethys to west Europe disappeared
temporarily (during the Eggenburgian and Ottnangian?). Recent discovery in Sihong
rendered it possible for us to compile a rather long list of forms common to both west
Europe and Asia. In addition to the two above cited forms, there are, for example,
Democricetodon, Megacricetodon, Amphicyon ... (vide supra). The sudden appearance
of so many common forms has demonstrated in a convincing way that rather free
faunal exchange between west Europe and Asia took place after the isolation period
lasting from middle Oligocene to early Miocene. The Indo-Pakistan subcontinent has
generally been considered as a separate zoogeographic region with highly endemic
fauna during the early Miocene, as evidenced by the Bugti fauna. It is interesting to
note here that some kind of similarity exists between it and the early Miocene faunas
of Gansu, China. For example, we can cite the following common forms: Sayimys,
Aprotodon, advanced forms of indricotheres, "Phyllotillon," and Gomphotherium.
Although the last two are widespread in Eurasia, the others have never been found in
west Europe. Therefore, the Indo-Pakistan subcontinent could possibly be considered
as a particular subregion of the Dinarid-Asian region.
Toward the end of the middle Miocene the faunal differentiation in separate
parts of the Eurasian continent became more and more clear. First of all, a number of
aberrant and endemic forms appeared in North China; for example, Gobicyon,
Platybelodon, Kubanochoerus, and "Oioceros." Gobicyon reached westward as far as
Yugoslavia; Platybelodon and Kubanochoerus were also found in the Mid-East and
Caucasus. Except for one mention (Tobien, 1973) of the probable presence of
Platybelodon in Sansan, no authentic record of its occurrence in west Europe has been
reported. "Oioceros" was probably restricted to central Asia. The Indo-Pakistan
subcontinent was characterized by the profusion of forest forms: Dryopithecus,
Sivapithecus and/or Ramapithecus, large number of anthracothere species, suids,
tragulids, bovids, etc., but no true cervids. There were also a number of forms
common to both Europe and Indo-Pakistan, but absent in central Asia; for example,
Deinotherium and Hyaenaelurus. So, faunistically the Indo-Pakistan subcontinent in
the middle Miocene might be closer to Europe than to central Asia. The famous coal-
bearing locality, Kaiyuan of Yunnan Province, where the first "Dryopithecus" in China
was found, should belong to the same paleozoogeographic province as the Indo-
Pakistan subcontinent, as judged by the revised faunal list (Dong W., 1987). Thus, the
boundary between the "Pro-oriental" and the "Pro-palaearctic" region during the late
middle Miocene can be drawn somewhere between Kaiyuan and Fangxian (a locality
with "Pro-palaearctic" fauna). Since no middle Miocene fossils have been found in
Tibet and other parts of South China, the boundary line must be rather arbitrary in
nature. What is certain is only that the boundary lies south to the present location,
which presently goes along the Qinling Mountains and the Huai River area.
Until the end of the Neogene this basic zoogeogrpahic pattern has remained
little changed. The "Pro-oriental" region gradually developed its present character.
However, the "Pro-palaeoarctic" region had undergone considerable change. Contrary
to west and central Europe, there was no clear indication of retention of forms
specially adapted to the forest habitat at the beginning of the so-called Hipparion
fauna. With the immigration of Hipparion from North America, central Asia seemed
to have immediately been occupied by a predominantly subtropical, dry and open
woodland fauna. This is the largest fauna both in number of forms and individuals, in
comparison with all the other faunas of the Cenozoic. In the latter half of the late
Miocene this fauna spread all over Eurasia. Perhaps, because of ecological reasons,
some forms were primarily confined to central Asia. The best known of this group was
the rhino Chilotherium, which probably never crossed the Aegean Sea (but see Guerin,
1980, p. 23), but was found in the Siwaliks. Some aberrant bovids, like Plesiaddax and
549
Urmiatherium, might also be restricted to Asia. Likewise, there were forms of
African origin that were absent in Asia, for example, Diceros, Orycteropus, and
Plioviverrops. During the late Miocene, Europe, central Asia, and the northern part of
Africa constituted a single zoogeographic region.
At the beginning of the Pliocene a new immigration event occurred; canids and
camelids crossed the Bering land bridge and entered Eurasia. Owing to the severe
desiccation of the Mediterranean Sea during the so-called Messinian crisis, these two
animal groups even penetrated the Iberian Peninsula. Thus, the faunal exchange
during the Pliocene was evidently more free than before. The same holds true for the
elephant and horse immigration events at the end of the Pliocene Epoch. They spread
rapidly all over Eurasia, including the Indo-Pakistan subcontinent. With deterioration
of temperature in the late Miocene, dry, subtropical, open woodland animals began to
vanish, or were driven to still more arid, even desert areas, while some others gradual-
ly adapted to temperate forest or shrub habitat, and thus survived. This eventually
made the dividing line between the north and the south zoogeographic regions more
distinct. The boundary line between the Palaearctic and Oriental regions must have
strongly fluctuated in accordance with the glacial cycles during the Quaternary.
From the foregoing discussion the following conclusions can be drawn:
First, during almost the whole Neogene (except for its very beginning) the major
part of the mainland of China had joined the rest of Eurasia to constitute a single
zoogeographic region. Several second-order provinces were gradually formed during
that time span. Because of the barrier provided by the Paratethys, west Europe was
separated several times from the rest of Eurasia, forming a special zoogeographic
province. During the second half of the late Miocene, west Europe was more isolated
than in the other periods.
Second, the Oriental region segmented from north Asia gradually. The "Pro-
oriental" region came into being probably no earlier than the second half of the late
Miocene. The middle Miocene fauna of the Yunnan Province was "Oriental" in nature,
and should be considered part of the Indo-Pakistan subcontinent zoogeographically.
Third, the faunal interrelationship between Eurasia and North America during
that time had been primarily limited in casual migration events. There were four
important dispersal events from North America to Eurasia: Anchitherium, l!ipparion,
Nyctereutes, Paracamelus, and Equus. From Africa there were two important disper-
sal events: Gomphotherium and Elephas. The above mentioned first appearance
datums are all extremely important in biochronologic calibration. There were other
dispersal events, but of only limited significance, because these immigrants did not
spread widely, often confined to limited areas, such as Diceros, etc.
NEOGENE MAMMAUAN FAUNA UNITS AND NOMENCLATURE PROBLEMS
After the faunal succession and its paleozoogeographic implication were dis-
cussed, the next step to be taken could be construction of a consistent Neogene bio-
stratigraphic scale and to fix it by using or introducing a suite of names of different
ranks as our European and American colleagues have been doing. However, here we
encounter some difficulties. First of all, the pursuit of Neogene biochronology in
China, taken as a whole, is still in its infancy. Probably the lack of important mineral
resources has attracted little attention to the Neogene continental deposits in China.
The mapping geologists were previously satisfied with the separation of large seg-
ments of time; e.g., Miocene, or Pliocene, irrespective of how poor the paleontologic
evidence. On the other hand, Neogene mammalian fossils were often found sporadi-
cally and identified without knowing their real superpositional relationships. The
faunas discussed under the Neogene Mammalian Fauna Succession of China section
represent only a series of disjunct samples in a line. Therefore, it seems the present
550
state of knowledge of Neogene mammalian faunas of China is too poor to meet the
basic demands for establishing formal biochronologic units. We refrain from calling
the above characterized faunas formal biozones. For the same reason we have not
established any formal biochronologic age either. However, we do feel it possible and
perhaps necessary to group these faunas into several informal biochronologic units
according to their developmental stages. These could well develop into formal
mammal ages of China in the future, provided they are better known and clearly
defined.
Another difficulty we met arises from the Chinese language. When the same
name is used to designate age-stage and fauna-formation, the way the people in the
western world distinguish the first from the second is simply to add an adjectival
suffix to the second. For example, in the languages of the western world, Barstow is
used to designate faunas and/or formation, while Barstovian designates a biochrono-
logic age. However, this cannot be applied in the Chinese language. In order to
distinguish them in the Chinese language we must add all the words of fauna, age ••• to
the proper name. Therefore, using the same name for fauna, formation, age, and
stage is a matter to be carefully considered, or perhaps to be avoided in Chinese (see
Yin z.x., 1979, p. 189). Otherwise, double meanings will inevitably appear for such
names. Unfortunately, the age names in the scale proposed by Li et al (1984) were all
taken from those of faunas. The present work shows that the majority of the faunal
names which were used by Li et al. to designate the ages, cover only a small part of
the time span of the given mammal ages bearing these names. As a result, almost for
all of such names, for example, Xiejia, Shanwang, Tung-gur (in Chinese), there are two
meanings (s.l. and s.s.). We noticed that our European colleagues tried to use regional
names to designate larger biochronologic units, for example, Agenian, Astaracian ••• ,
for stages or superstages. If we follow this practice, a series of new age-stage names
should be coined. We would like to leave this problem temporarily open. For the
moment we restrict ourselves to designate the faunal units only numerically.
Faunal unit I (see table Z) comprises the first three representative faunas,
Lanzhou, Xiejia, and Zhanjiaping plus the faunas thought to be contemporaneous with
them. We did not attach much importance to the probable first appearance of
proboscideans in Zhangjiaping, because, taken as a whole, the Zhanjiaping fauna is
characterized by retention of holdovers of Oligocene and possession of taxa closely
related to those of the Bugti fauna of Pakistan.
Faunal unit n comprises the Sihong and Shanwang fauna and localities attributed
to this unit. This unit is separated from the others by its close relationship with
Europe. A large number of animals are common with those of Europe, such as Demo-
cricetodon, Megacricetodon, Ursavus, Amphicyon, Plesiaceratherium, Dorcatherium,
and Lagomeryx.
Unit m is a period of profuse development of some rather endemic forms

typified by Platybelodon, Kubanochoerus, etc. These animals did not penetrate into
west Europe.
Unit IV is marked by the first appearance of Hipparion, along with the formula-
tion and the rapid development of the so-called Hipparion fauna, which spread all over
the vast land of Eurasia and Africa. In Europe a forest phase and a late savana phase
can be separated more or less clearly, but in China it is not so easy to separate the
Hipparion fauna into phases.
Unit V is faunistically distinct from the former unit in having newcomers from
North America (canids and camelids) and a large number of advanced forms derived
directly evolved from unit IV.
Unit VI is characterized by the immigration of Elephas (s.l.) and Equus in China

(see table Z).
551
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556
KEY BIOSTRATIGRAPmC EVENTS IN THE SIWALIK SEQUENCE
John C. Barry and Lawrence J. Flynn
Peabody Museum
Harvard University
Cambridge, Massachusetts 02.138, U.S.A.
INTRODUCTION
Miocene movements of crustal plates brought about fundamental changes in the

oceans by fragmenting the Tethys Seaway and further isolating the Atlantic, Indian,
and Pacific basins. Although controversial in the details of interpretation, the result-
ing paleoceanographic and tectonic events have been linked to global climate (Kennett
et al., 1985; Woodruff, 1985; Hodell et al., 1986) and, largely through their effects on
climate, can be seen to have had a far-reaching influence on the plants and animals of
the Miocene.
The same plate movements pushed the Afro-Arabian and Indo-Pacific plates
northward against Eurasia, creating chains of mountains and plateaux along the south-
ern margin of Eurasia and altering land connections among Africa, Eurasia, and the
Indian Subcontinent. In southern Asia the results of the collision of India and Asia
were particularly impressive, with the uplift of the Tibetan Plateau and the Himalayas
and other ranges. The same tectonic activity also produced linear zones of downwarp-
ing along the Asian-Indian suture, leading to the accumulation of the thick wedges of
sediment which now comprise the Paleogene and Neogene formations of India and
Pakistan. Most of the Paleogene is marine, although nonmarine parts in India and
Pakistan have produced important middle Eocene and older faunas. In contrast, the
Neogene is largely fluvial in origin and, loosely speaking, these continental Neogene
deposits are the Siwaliks.
The Siwaliks contain the best Neogene fossil record for southern and southeast
Asia and for that reason are of interest. But they also have an unusual fossil record,
one that encompasses long intervals of time with only minor hiatuses. This degree of
continuity is exceptional among terrestrial sequences and, since fossils occur through-
out the entire sequence, it provides a long, detailed record of biological and geological
events. This makes the faunas of the Siwaliks of special interest for understanding the
course of vertebrate evolution in southern Asia, the development of Eurasian terres-
trial environments, and the dynamics of Miocene faunal change in general. In this
paper we will discuss some of the major features of this record, highlighting biostra-
tigraphic events of interest and summarizing general conclusions of recent research.
GEOLOGICAL BACKGROUND
Oligocene and Miocene age sediments are found throughout Pakistan and India,
where they usually rest unconformably on middle Eocene or older deposits (Cheema et
al., 1977). In southern areas, as in the Kirthar and Laki ranges of Pakistan, Oligocene
marine rocks of the Nari Formation pass upward into a regressive sequence composed
of the Miocene Gaj and Man char Formations (Cheema et aL, 1977; Raz a et aL, 1984).
The age of the transition to continental sedimentation is not well established as,
except for the preliminary results of Khan et al. (1984), the formations have not yet
been systematically dated. However, the available paleomagnetic and paleontologic
evidence, which includes both the underlying marine faunas and the mammals, indi-
cates that the base of the terrestrial Manchar Formation is probably late early
Miocene (Cheema et al., 1977; Khan et aL, 1984; Raza et al., 1984; de Bruijn and
Hussain, 1984, 1985; Bernor et aL, 1988). To the north, in the Sulaimans and Kohat-
Potwar region, post-Eocene sediments appear to be entirely nonmarine, having been
deposited onto a land surface. The age of the onset of terrestrial sedimentation has
only been dated in the Salt Range of the Potwar Plateau, where it began just prior to
18 Ma (N.M. Johnson et al., 198Z, 1985). However, biostratigraphic evidence strongly
suggests that fluvial sedimentation may be progressively older to the south and west,
with a possible maximum age of late Oligocene/early Miocene (Raza and Meyer, 1984;
de Bruijn et aL, 1981).
Physical Stratigraphy
Exposures of the Siwaliks are very good on the Potwar Plateau and, because we
have been most involved in field and systematic research there, we will focus our
discussion on its sequence. The Potwar is also where most of the classic Siwalik litho-
and biostratigraphic units were first recognized (Pilgrim, 1910, 1913) and where the
most intensive recent geologic and paleontologic research has been conducted. There
the Miocene rocks are entirely fluvial in origin, having been deposited by la~ge rivers
and their tributaries. Total thickness of the deposits varies from north to south, but in
some sections over 3000 m accumulated and are now exposed at the surface. The
rocks are usually divided into time successive formations, with the classic sequence of
the Potwar comprising the Murree, Kamlial, Chinji, Nagri, and Dhok Pathan Forma-
tions of Pilgrim (1910, 1913) and what the Geological Survey of Pakistan refers to as
the Soan Formation (Cheema et al., 1977). All the lithological units are composed of
alternating sandstones and silts and clays, representing river channel and overbank
deposits. Individual formations have been distinguished by the ratio of sandstones to
siltstones, the maximum size of single sandstones, or their mineralogical or litholog-
ical composition (Cotter, 1933; Gill, 1951; Monaghan .in Pilbeam et al., 1979). It is
often difficult to delineate the boundaries between the formations, however, and from
the geological or sedimentological perspective it is best to view the Siwalik sequence
as a single genetic unit. Nevertheless, the Siwalik formations have always been
cryptic chronostratigraphic units and from the paleontological point of view recogni-
tion of the formations and their boundaries has been a crucial step in dating the fossils
(e.g. Colbert, 1935). This practice has in the past produced much confusion and sterile
debate, but we think it is now possible to assume that with the replacement of biostra-
tigraphy by magnetostratigraphy as a means of dating the rocks this era of confusion
is past.
Fossil sites are found throughout the entire sequence, although they are much
more common and richer at some levels than others. They are particularly productive
between 14 and 10.5 Ma and between 9.5 and 7.0 Ma and much of the recent collecting
has concentrated on these intervals. More recently, attention has shifted to the less
productive pre-14 Ma levels and knowledge of the older faunas is steadily improving,
although certainly not yet adequate. Significant recent discoveries include bovids and
tragulids that can be securely dated as being older than 18 Ma and we now have
several diverse assemblages of rodents from the interval between 18 and 14 Ma.
Efforts have also been directed toward the Pliocene and younger sediments, princi-
pally by Everett Lindsay and his students, but much remains to be done in the younger
levels.
Correlation and Age of the Siwaliks
Exposures of the Siwaliks on the Potwar and elsewhere are usually as large areas
of laterally contiguous outcrops. The largest of these areas are along the northern and
southern banks of the Soan River and along the northern flank of the Salt Range
558
between the Indus River and Jhelum. Exposures encompass tens to hundreds of square
kilometers and may be separated from other exposures by distances of over 50 km.
Within each area of outcrop it is usually possible to measure multiple stratigraphic
sections. By tracing marker beds laterally between them, a local lithostratigraphic
framework has been built up which, combined with a program of magnetostratigraphic
or radiogenic dating, provides a chronostratigraphic framework (Barry et al., 1980;
Tauxe and Opdyke, 1982).
Because of the disjunct occurrence of areas of outcrop and the highly variable
lithostratigraphy, correlations between areas can only be based on biostratigraphy or
magnetostratigraphy. The difficulties and opportunities of application of magneto-
stratigraphic techniques to terrestrial sequences are fairly well understood (e.g. Tauxe
and Opdyke, 1982; Lindsay et al., 1987). Some problems are technical in nature, such
as overprinting, weak magnetization, or inadequate sampling. They can usually be
overcome with appropriate laboratory or field protocols. Other difficulties are
geological or interpretive, such as the presence of unrecognized hiatuses or the neces-
sity of interpolating ages within magnetic polarity zones. They require careful obser-
vation in the field and cautious interpretation of results (Badgley et al., 1986). Never-
theless, with application of magnetostratigraphy on a large scale, it has proved
possible to construct a single composite Potwar sequence and successfully correlate it
to the geomagnetic time scale (Opdyke et al., 1979; N.M. Johnson et al., 1982, 1985;
Tauxe and Opdyke, 1982). Probably the most important result of this effort has been
to establish that the Potwar Siwaliks span over 17 million years without any major
hiatuses, although several local sections have significant breaks and the interval
between 6 and 3 Ma is not well represented by exposures. The base of the Miocene is
widely exposed throughout the Potwar and the oldest date yet obtained is from a
section on the northern flank of the Salt Range near Chinji village. There the base is
more than 18 Ma (N.M. Johnson et al., 1985). Very young rocks are more restricted in
occurrence, but the 700,000 year old Brunhes-Matuyama boundary has been recognized
in sections on the eastern end of the Salt Range near Jhelum (Opdyke et al., 1979).
The magnetostratigraphic correlations agree with fission track ages on late Pliocene
and late Miocene bentonites (G.D. Johnson et al., 1982).
Single fossil sites can be traced laterally into the local stratigraphic sections, so
that their relative stratigraphic positions can be established solely on the basis of
superposition. As a result we can now establish the ages of a great many of our local-
ities to within 200,000 or even 100,000 years; exceptional precision for a terrestrial
sequence.
Taphonomy, CHmate, and Physical Environment
The vertebrate fossils have two modes of occurrence; as rich but small and local
concentrations or as an ubiquitous scatter of isolated specimens. The abundance of
sites and fossils varies, but all formations have at least occasional poorly preserved
bones and teeth and usually small concentrations of identifiable material can be
discovered as well. Taphonomic and sedimentological studies show that the fossils
occur predominantly in fluvial environments, with concentrations being found most
frequently in the main and secondary channels, in levees, and in the flood deposits of
the smaller streams (Badgley, 1982, 1986; Badgley and Behrensmeyer, 1980;
Behrensmeyer, 1987). We have also found fossils in the paleosols of the ancient flood
plains, in small ponds, and, as predators were also important in concentrating bones,
very occasionally in burrows, dens, and scat or pellet concentrations (Badgley, 1986;
Behrensmeyer, 1987). The latter type of deposit is particularly important for rodents
and we have found them to be much more common than had been previously sup-
posed. The richest concentrations typically are sites that include more than one of
these general depositional environments. Most sites formed over periods ranging from
tens to thousands of years, implying that single localities might contain animals that
lived several thousand years apart and perhaps in different habitats and separate
ecological communities. Such mixed assemblages are disadvantageous for recon-
structing the original communities at the most local level. However, time-averaging
and community mixing do serve to homogenize individual sites and by removing unique
559
features make them more comparable to each other. This is particularly important
when comparing sites separated by millions of years or from different geographic
areas and it has a critical influence on biostratigraphic correlations.
A synthesis of our taphonomic, sedimentological, and paleontological results

forms the basis of preliminary inferences about Siwalik climates and possible habitats
of Siwalik faunas. Middle and late Miocene climates of northern India and Pakistan
were probably monsoonal, as at present. Otherwise, little is known about them beyond
what can be inferred from the floral record and from oceanographic data. On the
Potwar Plateau fossil plants associated with vertebrates are rare (although not
unknown), but paleobotanical studies of contemporaneous Indian deposits (Lakhanpal,
1966; Prasad, 1971; Prakash, 1973; Mathur, 1984) suggest the presence by the early
Miocene of a wide variety of plant communities and diverse habitats. Deciduous
species are present in the early Miocene sites, as are grasses, but there is little
evidence for a progressive change to drier or more seasonal vegetation until later in
the Miocene. These conclusions, however, are based on fossil pollen and macrofloras
from the east and south in India, a region which may have had a somewaht different
climate than the Potwar in the middle Miocene. A more humid climate than at
present, nevertheless, is in accord with the diversity of vertebrates and the types of
sediments and paleosols f~und on the Potwar. With the northward movement of India
and late Miocene intensification of atmospheric circulation and aridification, the
Potwar climate would have become progressively less humid and more seasonal. This
may have culminated in a subhumid climate at the end of the Miocene and perhaps a
semiarid climate during periods of the late Pliocene and Pleistocene.
During the early and middle Miocene the Potwar was proably of low relief, with
one or more very large rivers and many smaller ones. The river flood plains must have
been extensive and undoubtedly contained a wide variety of habitats. There seem to
have been few permanent lakes or ponds, but low areas may have held seasonal
swamps and ephemeral oxbow lakes would have formed in abandoned stream
channels. The ecological reconstructions of the lower and middle Siwalik faunas
suggest primarily woodland or even forested habitats (Andrews, 1983; Kappelman,
1986). However, near larger streams the vegetation should have been continually
disturbed by flooding and this could have left a mosaic of small areas of brush and
grassland intermixed with more closed forest.
Global Climate Events
The changes in diversity and faunal composition of Siwalik terrestrial verte-

brates show general, but weak, correlations to global climate and oceanographic
events, which are reviewed in the following. While early Miocene climates appear to
have been relatively equable and stable, several lines of evidence indicate that
between ca. 16 to 1Z Ma there were substantial changes in the physical environment of
the earth. Intensification of oceanic and atmospheric circulation, steepening of
temperature gradients, and expansion of Antarctic glaciation are particularly note-
worthy (Kennett et al., 1985; Kennett, 1986; Savin et al., 1985; Woodruff et al., 1981;
Woodruff, 1985). In addition there were major oscillations in eustatic sea level near
16 Ma and two smaller falls between 14 and 1Z Ma (Haq et al., 1987).
Beginning at about 1Z Ma and continuing to about 8 Ma isotopic studies suggest

temperatures were variable, but without significant trend (e.g. Kennett, 1986; Savin et
al., 1985; Woodruff et al., 1981). However, two cool episodes bracketing the middle to
late Miocene boundary have been identified. The biostratigraphic and paleomagnetic
correlations indicate an age between 11.0 and 9.5 Ma for the older episode, while
those given for the younger suggest it is probably between 9.0 and 8.0 Ma (Kennett,
1986; Berggren et al., 1985). The second episode is one of the largest isotopic excur-
sions in the Miocene (Kennett, 1986). In the same middle and late Miocene interval,
the Haq et al. (1987) eustatic curve has a progressive fall in sea level ending at about
10.5 Ma with one of the lowest stands of the entire Neogene. Keller and Barron (1983)
and Barron et al. (1985) have also identified middle Miocene deep sea hiatuses and
correlated them to isotopic events. Their NH4 and NH5 hiatuses are interpreted as
560
due to climate change and reorganization of oceanic circulation. The correlation and
dating of these various events is necessarily uncertain. The middle to late Miocene
low sea level, the first of the isotopic excursions, and the NH4 hiatus may all record a
single, major global climatic event at ca. 10,5 Ma.
The rest of the late Miocene (ca. 8-5.3 Ma) had at first somewhat warmer condi-
tions (Kennett, 1986), with very marked cooling after 6.5 Ma. The cooling trend is
present in the faunal and oxygen isotope records (e.g. Burckle, 1985; Kennett, 1986)
and can be related to a prominent carbon isotope shift and the eventual desiccation of
the Mediterranean (Vincent et al, 1985; Hodell et al., 1986). Latest Miocene deep sea
hiatuses and sea level oscillations have also been noted (Keller and Barron, 1983; Haq
et al., 1987). These paleoceanographic events are probably related to the widespread
increasing seasonality and aridity of terrestrial environments in the late Miocene
(Jacobs and Flynn, 1981; Flynn and Jacobs, 198Z; Janecek and Rea, 1983; Stein, 1985;
Van Zinderen Bakker and Mercer, 1986).
BIOSTRATIGRAPHY
Siwalik Biostratigraphic Zonation
The history and usage of biostratigraphic terminology in the Siwaliks is one of

confusion. Pilgrim (1910, 1913) first recognized a series of successive "faunal zones,"
initially using the term in a manner comparable to modern usage of the "stage"
concept. Pilgrim's units (Kamlial, Chinji, etc.) were based on a mixture of contained
fauna and lithological criteria. In most instances their superpositional relationships
could be demonstrated, but the boundaries of the faunal zones were not delineated
and, because of mistakes in correlation, the faunal content of some zones could never
be adequately differentiated. Subsequently, as stratigraphic concepts and nomencla-
ture became more precise, Pilgrim's faunal zones came to be used primarily as litho-
stratigraphic formations, as chronostratigraphic "zones," or some confused combina-
tion of both (Pilbeam et al, 1979; Barry et al., 1980), Only occasionally were they
recognized as being essentially biostratigraphic units. Because of this change in usage
and the resulting confusion, Barry et al. (1980, 198Z) first advocated restricting
Pilgrim's terms to the lithostratigraphic formations and later proposed a new series of
biostratigraphic zones to replace the middle and upper Siwalik "faunal zones." The
intention was to eventually define zones for all the formations and, if feasible, sub-
divide the existing zones. Principally for that reason, the zones were formulated as
interval-zones. That is, each zone had a defined base and included all of the strati-
graphic interval below the base of the succeeding zone. The base of each zone was
defined by an evolutionary event and for ease of recognition first appearances of
abundant and easily identified immigrant species were selected as the events. How-
ever, other types of events, including extinction or speciation events, and other types
of biostratigraphic zones, especially assemblage or concurrent-range zones, could have
been used as well. All could be combined into one hybrid zonal scheme.
Murphy (1977) has noted the distinctions between the operations of definition,
characterization, and identification in the practice of stratigraphy. The biostrati-
graphic interval-zones of Barry et al. (198Z) were defined and characterized in strati-
graphfc reference sections and criteria were stated for identifying or recognizing
them in other sections. Because they were related directly to stratigraphic sections,
the interval-zones and their boundaries can, like stages, be correlated to other geolog-
ical phenomena, such as sedimentological or geochemical events, magnetopolarity
zones, or to geologic time. However, the Siwalik interval-zones should not be con-
fused with stages, which are chronostratigraphic units. Each interval-zone's lower
boundary is defined by a biological event, not a stratigraphic level having a specific
age. At the time the defining taxa were selected, the stratigraphic levels and ages of
their first appearances were thought to be accurately known. However, unlike stages
(Ager, 1973), because the Siwalik interval-zone boundaries are based on biological
events, they are not fixed in the reference sections or in time. If subsequent collect-
ing were to demonstrate an older occurrence for a defining taxon, then the boundary
561
of the corresponding biostratigraphic unit would also be adjusted downward in the
reference section.
This formulation of the zones may trouble some readers, but the Siwalik bio-
stratigraphic interval-zones as proposed are not chronostratigraphic units. The
Siwalik faunas appear to belong to a distinctive zoogeographical province (Bernor,
1983) and we think the biostratigraphic interval-zones may eventually prove to be
useful as chronostratigraphic zones, but that transformation is an independent step.
Their utility as chronostratigraphic zones depends on how consistent their temporal
relationships are in other sections. If the events that define the biostratigraphic
interval-zones prove to be isochronous throughout the Siwalik faunal province, then
they will also define chronostratigraphic interval-zones, but ideally their isochrony
should be demonstrated, not assumed. In most situations, the isochrony of chrono-
stratigraphic zones can only be tested by the weak method of finding internal inconsis-
tencies, such as inverted zones or extreme difficulty in identifying a zone. In the
Siwaliks, however, the widespread application of magnetostratigraphy makes it possi-
ble to test boundary isochroneity by determining how consistent their stratigraphic
relationships are to magnetozones in numerous sections.
Therefore, in current usage, Pilgrim's terms are applied only as lithostrati-

graphic divisions. The biostratigraphic framework is independent and, as it matures
with ongoing collecting and improved stratigraphic control, the ages of the interval-
zone boundaries will become more precisely known. Once their isochroneity is tested
in other sections, they will form the basis for chronostratigraphic units.
Fauoal Events
Although previous systematic work on Siwalik vertebrates had already shown

them to be diverse, new and more intensive collecting techniques have recently added
many taxa, especially of smaller species such as rodents, lagomorpha, primates, bats,
and insectivores. The new collections have also extended the ranges of others. As a
consequence, the fossil assemblages are more balanced ecologically and the strati-
graphic ranges of many species are known with greater precision. Another particu-
larly relevant conclusion of recent systematic revisions is recognition that the large
mammals are neither as diverse nor geographically distinctive as had been thought.
We find many groups have some connections with extant Southeast Asian mammals
and Siwalik and African middle Miocene faunas especially are similar. There are,
however, differences; a puzzling example of which is the co-occurrence of cercopithe-
coids and hominoids in East Africa and their temporal separation in the Siwaliks. This
suggests something of the zoogeographic and temporal complexity of mammal evolu-
tion during the Miocene.
First and last occurrences of many Siwalik mammals can be recognized using
recently developed stratigraphic range data and in Table 1 we present an updated list
of 133 events. These events approximate the local appearance and local extinction of
Siwalik species and are of interest as evidence bearing on questions concerning tempo
and mode of speciation and the formation and evolution of the vertebrate communi-
ties. In the present context, they are also of interest because they are a basis for
biostratigraphic correlations to European and other sequences. In all cases the ages
listed in Table 1 are for the oldest or youngest specimen in one or more :iotwar
sections and are based on estimates derived from paleomagnetic correlations to the
geomagnetic time scale of Mankinen and Dalrymple (1979).
As estimates of local first appearances and extinctions, the events in Table 1

differ in precision of resolution. For some taxa and stratigraphic intervals the ages
may be accurate to within 100,000 years. Other ages are not as reliable, but we have
not attempted to distinguish between those we have confidence in and those we do
not. It is, however, especially difficult to establish first or last occurrences for
uncommon or rare species and they are the majority in Table 1. While attempting to
present as many events as possible, we have been selective in which events we have
chosen to present. It is inevitable that with additional collecting and systematic study
562
the dates will shift, but the uncertainty of some is compensated for by their biostrati-
graphic interest. (It may be of some interest to compare Table 1 to previous compila-
tions: Opdyke et al., 1979; Jacobs and Flynn, 1981; Flynn and Jacobs, 198Z; Barry et
al., 198Z, 1985.) We have the greatest confidence in ages of large species that are
easily identified and common. These include Stegodon, Elephas, the hipparionines,
Eguus, Hexaprotodon, Listriodon, Conohyus, Giraffa, and the large bovid Selenoportax
lydekkeri. Sites with small mammals are less common and small mammal ranges are
q~ore likely subject to errors on the order of a half million years or less, depending on
the quality and density of samples. Otherwise, because good sites tend to have very
diverse small mammal assemblages, we have a greater degree of confidence for them
than for most of the larger mammals.
Badgley and Gingerich (1988) have discussed how the apparent patterns of first
and last occurrences used to define faunal turnovers can be an artifact of sampling
and varying fossil productivity among stratigraphic levels. They note that maxima of
appearances coincide with large samples from highly productive horizons, while
maxima of disappearances coincide with small samples from poorly productive
horizons; an effect that can distort the apparent ages of the events. Since there is
considerable variation of fossil productivity among levels in the Siwaliks, this effect is
also important there and must be considered as a limitation on interpretations of the
record. Figure 1 is a representation of our estimates of the quality of data (or produc-
tivity) for the rodents and artiodactyls in the most critical middle and late Miocene
part of the Potwar sequence. We have divided the time from 16 to 7 Ma into nineteen
equal intervals. Because the mode of origin and methods of collecting are different
for small and large mammals, we have assessed these groups separately. Quality is
expressed as a value between 1 (best quality) and 5 (no data) and has been determined
from the number of specimens collected, the number of sites, and the preservational
quality of the fossils in the interval. These estimates are based on the rodents and
artiodactyls, but can be assumed to roughly reflect the quality of other small and
large mammals as well.
Figure 1 shows that the best quality data comes from the 16, 14-13.5, 11-10.5,
9.5-9.0, and 8.0-7.0 Ma intervals (average quality Z.5 or better). These intervals would
Rodent•
2
3
7 8 9 10 11 12 13 14 15 18
Artlodectyla
2
3
4
5 ~~~~~~~~~~~~~~~~~~~~~~
7 8 9 10 11 12 13 14 15 16
Million• of Yeera
Fig. 1. Quality of data for each half million year interval between
16 and 7 Ma inclusive. Class 1 =best quality, Class 5 =no
data.
563
Table 1. Key Biostratigraphic Events in the Siwalik Sequences
> indicates undoubtedly or likely to be older than; ? indicates identification is

uncertain; ca. indicates age is approximate.
First Appearance Last Appearance

(Ma) (Ma)
Sciuridae > 18.0

Gliridae 13.5 7.0
Sayimys spp. > 18.0 8.8
Thryonomyidae > 16.1 1Z.7
Copemyinae > 18.0 8.8
Copemyine reappearance 7.6 7.5
Megacricetodontinae > 16.1 10.6
Myocricetodontinae > 16.1 1Z. 7
Dendromurinae > 16.1 9.4
Protatera sp. 7.6 7.6
Prokanisamys benjavuni > 16.1 ca. 11.6
Kanisamys indicus > 16.1 10.6
Kanisamys potwarensis 13.5 1Z.7
Kanisamys nagrii 10.8 8.8
Kanisamys sivalensis 8.5 ca. 7.0
Protachyoryctes tatroti 6.9 ca. 6.5
Eicooryctes kaulialensis 6.9 ca. 6.4
Rhizomyides sivalensis ca. 6.4 ca. 5.5
Rhizomyides punjabiensis 9.4 9.Z
Rhizomyinae spp. 8.5 ca. 7.0
Anepsirhizomys opdykei ca. 3.0
Antemus chinjiensis 14.3 11.8
Mus.!:!!£12!:. 5.5
Hadromys loujacobsi 1.5
Progonomys spp. ca. 11.6 ca. 7.0
Karnimata spp. ? 10.6 5.5
Parapodemus sp. 8.1
Parapelomys robertsi 5.5
Golunda kelleri 1.5
Hystrix sivalensis 7.0
Leporidae 7.0
Lorisidae ? 16.1 ca. 7.0
?Presbytis sivalensis 6.3
Sivapithecus spp. ca. 1Z.O 7.4
Gigantopithecus bilaspurensis 6.3
Dionysopithecus sp. 16.1
Metapterodon n. sp. 8.8
Dissopsalis carnifex > 16.1 8.8
Amphicyonidae > 18.0 ca. 7.0
Indarctos punjabiensis ca. 6.5
Herpestes spp. ? 1Z.7
Hyaenidae 13.Z
564
First Appearance Last Appearance
(Ma) (Ma)
Percrocuta carnifex ? 1Z.6

Percrocuta grandis ca. 5.5
Nimravidae > 15.1 7.4
Deinotheriidae > 18.0 7.0
Elephantoidea > 18.0
Stegodon ca. 4.5
Elephas planifrons Z.9
"Hipparion" spp. 9.5 1.5
Equus sivalensis Z.5
Chalicotherium salinum 6.5
gigantic anthracotheres ca. 18.0
Hexaprotodon sivalensis 5.3
Sanitherium schlagentweit i > 16.1 11.8
Listriodon spp. > 16.1 9.4
Conohyus spp. > 15.0 9.4
Tetraconodon magnus 9.0 8.3
Sivachoerus prior Z.9
Hippopotamodon sivalensis 9.Z 6.6
cf. Propotomochoerus hysudricus 11.3 ca. 5.5
Potamochoerus sp. 4.0
Sus sp. Z.9
cf. Kolpochoerus sp. Z.9
Hippohyus sivalensis Z.9
Dorcabune spp. > 16.1 7.5
Dorcatherium spp. > 18.0 6.4
cf. Dorcatherium sp. 4.0 Z.9
large giraffoid > 18.0 ca. 16.0
Giraffokeryx punjabiensis ca. 16.0
?Bramatherium megacephalum 6.6
Sivatherium giganteum Z.9
Giraffa punjabiensis 7.3
Cervidae Z.5
?Eotragus sp. A > 18.0
?Eotragus spp. 15.Z 1Z.8
Kubanotragus sokolovi 13.8 11.4
Sivoreas eremita 13.8 10.6
Helicoportax tragelaphoides 13.0 11.4
Protragocerusgluten 13.8 10.6
Sivaceros gradiens 13.8 10.6
Tragoceridus pilgrimi 9.0 8.1
Kobus sp. ca. 7.0 ca. 3.0
Selenoportax lydekkeri 7.4 ca. 4.0
Proamphibos lachrymans ca. 4.5
133 events
565
then be expected to have large numbers of first appearances, while the succeeding
intervals should have the most disappearances. This prediction seems to be borne out
by Table 1 and this consideration implies that we must exercise cautious judgment in
interpreting the significance of any dates in Table 1. Rare taxa, such as the
hominoids, or those which can only be identified from limited material, such as the
bovids, are especially vulnerable.
If an interval has abundant fossils and many sites, it is reasonable to infer that
failure to find an otherwise common species indicates its true absence, rather than
being an artifact of incomplete sampling. However, when an interval has few fossils,
it is much more difficult to establish that a taxon is absent. Nevertheless, for species
that are very common within their demonstrated stratigraphic range, it may still be
possible to establish their absence. Whether a species is common or rare, the
approach depends on establishing an expectation of finding it and then determining
whether the expectation is fulfilled or not (Nichols and Pollock, 1983). In the case of
the first appearance of hipparionines (at a site older than the one discussed by Badgley
et al., 1986), the immediately underlying stratigraphic levels have few fossils (Fig.
1). Nevertheless, bovids, giraffes, tragulids, and various suids have been found in
these levels. Within their demonstrated range zone, fossil remains of hipparionine
horses are normally much more common than those of any other taxa and we would
therefore expect to find at least some fragmentary hipparionine material. Since we
have not, despite diligent searching, we have tentatively concluded that equids were
not present in the Siwaliks before approximately 9.5 Ma.
A final caution concerns the dates listed in Table 1. As previously noted, these
are derived from correlations to the geomagnetic time scale of Mankinen and
Dalrymple (1979). Alternative scales have been proposed, however, and these assign
different ages to most of the middle and late Miocene magnetic chrons. On the widely
used Berggren et al. (1985) time scale the ages of the C5 Chrons are as much as
400,000 years older than on the Mankinen and Dalrymple (1979) scale.
FAUNAL CHANGE
An overview of the nearly eighteen million years of faunal evolution documented

on the Potwar highlights three major phases of change. The first encompasses the
establishment of the Siwalik faunas prior to 18 Ma and their initial phase of develop-
ment prior to 14 Ma. During this phase bovids and other ruminants became the most
diverse and ecologically dominant large herbivores, a position they still hold in the
modern world. An earlier ctenodactyloid rodent fauna was replaced by a small
mammal fauna of mainly advanced muroid rodents; early and middle Miocene assem-
blages were dominated in both numbers and diversity by cricetids. Although their
record is much less complete, it is very likely that the aeluroid carnivores underwent a
parallel radiation.
During the second phase, between 14 and 9.5 Ma, there were episodes of faunal
change, but these do not appear to have led to any fundamental community reorgani-
zation. Included are the diversification of bovids and hyaenids, appearance of large
hominoids, and origin of murids. At 9.5 Ma hipparionine horses appear and there is a
brief interval with extinction of older species, notably suoids and cricetids. Murids
are more dominant than cricetids by this time, both in diversity and abundance.
Whether the turnover at 9.5 Ma also involves a major ecological reorganization is an
unresolved problem. The immigrant taxa in themselves probably had a major ecolog-
ical impact, but at present we are not able to demonstrate that pre-hipparionine
habitats were much different from later ones.
Between 7.5 and 6.5 Ma the Siwalik fauna for the first time becomes closely
similar to contemporary ones in northern and western Eurasia. This pattern of
Eurasian connections contrasts to previous similarities to modern Southeast Asian
mammals and indicates a new phase had begun. Extinctions include Sivapithecus,
archaic carnivores, Deinotherium, Brachypotherium, dormice, and possibly tree shrews
566
and lorisids. Appearances include leporids, Hystrix, a giraffine, several bovids, and
eventually cercopithecids.
In the Siwaliks first and last records occur in every interval, but also form
discrete clusters of above normal faunal change. This clustering gives the record an
episodic aspect. At present we recognize seven episodes of faunal change, which are
dated at older than 18 Ma, 14-13.5 Ma, 12. Ma, 9.5 Ma, 7.4 Ma, 5.3 Ma, and 2..9 Ma
(Opdyke et al., 1979; Barry et al., 1982., 1985). Maxima in first and last occurrences
may coincide, as between 8 and 7 Ma, or they may be disjunct, as with the maximum
of appearances at 14-13.5 Ma and maximum of disappearances at 12..0 Ma. During
turnovers faunal change is the result of in situ evolution, immigration, and extinction,
but the relative importance of these three processes is uncertain. In the Siwaliks in
situ evolution appears to occur in only a few lineages and is therefore thought to be
unimportant in most groups. Immigration and extinction events tend to be correlated
and together were the principal cause of faunal change. As immigration events often
precede extinctions, and in some cases can be inferred to have caused them, immigra-
tion and the resulting ecological disruption may have been the primary cause of
community change. Many of the immigrant species probably originated in Africa,
Europe, or other parts of Asia. Faunal turnovers are thus also intervals of faunal
exchange and indicate times when land connections were established. Nearly all these
episodes show approximate correlations to global climatic, oceanographic, and
tectonic events, and these, through their effects on sea-level, intercontinental
connections, and vegetation, may have controlled movement of mammals into the
Siwalik province. Separating these factors would be difficult, however, and the time
resolution of these environmental and faunal events is too poor to conclusively link
them. Nevertheless, the distinct clustering of events prior to 7.4 Ma suggests that at
that time discrete episodic environmental events, such as sea-level changes, were
most influential. After 7.4 Ma faunal change is more diffuse; possibly because
climatic change had become more important.
The Siwalik faunas had their origin in a major turnover which occurred prior to
18 Ma and possibly as early as 2.2. Ma. That turnover event is manifest by the appear-
ance of many immigrant taxa in the Siwaliks (Barry et al., 1985) and can be approxi-
mately correlated to oceanographic events, including a short term fall in sea-level
(Haq et al., 1987). As a consequence, the middle and late Miocene Siwalik faunas are
a mixture of endemic Asian taxa and immigrant-derived taxa. After 18 Ma, there are
several turnover events, some of which are more abrupt and of greater magnitude than
others. However, throughout its history, and particularly just prior to 7.5 Ma, the
Siwalik fauna continues to have a strong taxonomic similarity to the modern fauna of
tropical Southeast Asia and it is likely that many extant Southeast Asian species are
derived from or closely related to Siwalik forms.
The taxonomic composition of the post-7 .0 Ma fauna suggests a major ecological

change took place between 7.5 and 7.0 Ma. This is of particular interest because the
large hominoids probably became extinct at 7.4 Ma, while cercopithecoids make their
first appearance shortly afterward. Sivapithecus presumably occupied closed habitats
and its local extinction and that of many other species after 7.4 Ma seems to point to
more open or more seasonal latest Miocene Siwalik habitats. At the present time, we
link this change to increasing aridity (Flynn and Jacobs, 1982.; Jacobs and Flynn,
1981). The increase in seasonality, however, does not seem to have been widespread,
as contemporaneous faunas only 300 km to the southeast at Haritalyangar retain a
distinctly tropical woodland aspect. We would expect this, however if the Potwar
faunal change was the result of the eastward extension, or intensification, of a sub-
tropical latitudinal arid zone, combined with restriction of the strong monsoonal
influence to more easterly regions of the subcontinent.
Data on species durations, the degree of overlap between closely related species
(or other pairs of potential competitors), and the rate and tempo of morphological
change within individual lineages are available for some Siwalik mammals. Most
species are rather short--lived. For example, rodent species longevities average about
Z to 3 million years. But some species may last very much longer, with, for example,
567
ranges of rhinoceroses approaching 10 million years (Heissig, 1972). Preliminary
analysis (Flynn, 1986) also suggests that in a few clades average species durations
decreased significantly in the late Miocene as compared to the middle Miocene. This
is possibly due to increasing frequency of environmental disturbances, since Siwalik
extinctions have some correlations with global environmental events (Barry et al.,
1985) and environmental events are an important cause of extinction and speciation,
as noted by Vrba (1985), Webb (1984), and others. In this context the period between
17 and 13 Ma should be of special interest, for it is an interval of rapid and irregular
climate change and it is also when ruminants, advanced muroids, and aeluroid carni-
vores diversified to become the dominant mammals of the Old World tropics. If the
frequency of environmental disturbances is directly related to species longevity and
turnover, then we would expect that between 17 and 13 Ma species durations should be
shorter than average and turnover more rapid. Possible connections between species
longevities, rates of turnover, and the increasing diversity of muroids, ruminants, and
carnivores remain an intriguing puzzle.
ACKNOWLEDGMENTS
The efforts of many individuals have provided the information used in this
survey. We particularly wish to thank William Downs, Everett Lindsay, Louis Jacobs,
Mahmood Raza, and Nikos Solounias for identifications and David Pilbeam for
support. Funding for the research has been through the National Science Foundation
and the Smithsonian Institution's Foreign Currency Program.
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Prasad, K.N., 1971, Ecology of the fossil Hominoidea from the Siwaliks of India.
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570
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571
QUO VADIS, ANTEMUS? THE SIWALIK MUROID RECORD
Louis L. Jacobs
Shuler Museum of Paleontology

Southern Methodist University
Dallas, Texas 75275, U.S.A.
Lawrence J. Flynn
Peabody Museum
Harvard University
Cambridge, Massachusetts 02138, U.S.A.
William R. Downs
Shuler Museum of Paleontology

Southern Methodist University
Dallas, Texas 75275, U.S.A.
and
Bilby Research Center
Northern Arizona University
Flagstaff, Arizona 86001, U.S.A.
John C. Barry
Peabody Museum
Harvard University
Cambridge, Massachusetts 02138, U.S.A.
INTRODUCTION
The fossil rodent Antemus is the oldest member of the most diverse family of
extant mammals. It is known from the Miocene of southern Asia as part of a distinc-
tive evolving fauna with complex biogeographic relationships to Europe, Africa, and
other parts of Asia. The Neogene small mammal record in southern Asia, of which
Antemus is a part, is dominated by muroid rodents. The Siwalik section of the Potwar
Plateau, northern Pakistan, provides most of the available fossil evidence for the
history of southern Asian rodents.
In the following, we discuss the biostratigraphy of southern Asian rodents as it

applies to European Neogene chronology. We also examine the pattern of first occur-
rences, last occurrences, and relative abundance in the Siwalik muroid record in order
to distinguish major events in the Neogene history of rodents in southern Asia.
The Potwar Siwaliks embody an essentially continuous, if episodic, record of

Miocene mammals from 18.3 to 5.5 Ma. There is a Pleistocene assemblage as well
(Musser, 1987), and potential for development of the Pliocene record (Opdyke et al.,
1979). The sequence of fossil sites is well calibrated temporally (e.g., Barry et al.,

--- l
I
I
I
I
I
.I
( "
PAKISTAN
- 1 00 km
eBUGTI
GAJ
•
• N
i
Fig. 1. Map of Pakistan showing the major collect-
ing areas discussed in the text. Gaj and
Sehwan are sections through the Manchar
Formation; Banda Daud Shah includes both
Murree and Chinji Formation sites. Daud
Khel is about 60 km southeast of Banda
Daud Shah.
1985). Other important small mammal assemblages from elsewhere in Pakistan extend
the record of the early Miocene. These are primarily Bugti in Baluchistan (Jacobs et
al., 1981; Flynn et al., 1986), the Manchar sections of Sind (de Bruijn and Hussain,
1984), and the Murree and Chinji Formations west of the Potwar (de Bruijn et al.,
1981; Wessels et al., 198Z; Munthe, 1980; see figures 1 and Z).
Dating of origination and dispersal events determined in the Siwalik section can
provide limits for biogeographical and biochronological scenarios in Europe. The
systematics of circum-Mediterranean muroids is relatively well known (for close
relatives of Siwalik taxa, see Michaux, 1971; Fahlbusch, 1964; Mein and Freudenthal,
1971; Engesser, 197Z; Jaeger, 1977; Chaline and Mein, 1979), but the absolute chronol-
ogy of European species is generally uncertain. A case in point is the earliest occur-
rence of Progonomys. In Europe it is close to (perhaps synchronous with) the
"Hipparion datum" and the commencement of the Vallesian (MN 9). Dating of the first
appearance of Hipparion in Europe is controversial. Hipparionines in Pakistan first
occur at approximately 9.5 Ma, while Progonomys occurs significantly earlier.
Refinement of Siwalik Progonomys chronology, presented below, provides an addi-
tional line of evidence bearing on European biochronology and thereby tests the bio-
geographic significance of the "Hipparion datum."
574
p'oTWAR BANDA DAUD SHAH,
GAJ SEHWAN
BUGTI
Ma <
<
12
CHINJIFM.
<
I I
I I H-GSP 8224
<
: H-GSP 107 I H-GSP 8227
14
~
I
~///////////, M~F~ ~F~
<
KAMUALFM.
16 < Y592 T1H-GSP 8114, 52

I
I H-GSP 8114A
I
16 < Y721
I H-GSP 8106 ~//////////h
I
H-GSP 116 ..!...
GAJFM.
20
GAJFM.
22 BUGTI
Fig. z. Provincial correlations within Pakistan (modified from

Bernor et al., 1988). On the left are the Potwar lower
Siwaliks with paleomagnetically dated rodent sites (indi-
cated by chevrons). Sites YGSP 59Z and YGSP 7Z1 are
crucial in showing that H-GSP 116 antedates 18 Ma.
Dashed vertical bars indicate age range limits constrained
by Potwar localities; the lower limit for H-GSP 8106 is
H-GSP 116.
POTWAR BIOSTRATIGRAPHY
The Siwalik rock sequence has been dated paleomagnetically in a series of

studies (Johnson et al., 1985; Keller et al., 1977; Opdyke et al., 1979; Tauxe and
Opdyke, 198Z) and spans 18.3 Ma to less than 1 Ma, where dated. In the Potwar
Plateau, rocks are well exposed and readily traced laterally. The middle and late
Miocene are particularly well represented by rodent localities that can be shown to be
time successive by superposition. Thus, stratigraphic relationships are not in question
for the local area of the Potwar Plateau. Absolute ages are interpolated relative to
magnetic reversal events. While a particular absolute age assignment for a site may
be inaccurate on the order of 100,000's of years, depending on choice of time scale
used (here, Mankinnen and Dalrymple, 1979), or on varying rates of deposition, careful
stratigraphic work allows unambiguous ordering of sites.
The biostratigraphy that results from plotting observed temporal distributions of

species permits direct correlation within the South Asian biogeographic province, e.g.,
with the Manchar Formation of southern Pakistan (see figure Z, altered from Bernor et
al., 1988). Rodents from H-GSP 8106, particularly species of Sayimys and the muroids
Prokanisamys and Spanocricetodon, are primitive with respect to those from low in
575
the Potwar section, suggesting an age in excess of 18 Ma. The base of the Murree
Formation at Banda Daud Shah is older still, as indicated by the rodent fauna (de
Bruijn et al., 1981; de Bruijn and Hussain, 1984). H-GSP 116 may predate significantly
the base of the section on the Potwar Plateau, where it is dated at 18.3 Ma. Although
difficult to correlate individually, sites H-GSP 116, 8106, and 8114A are a temporally
successive series (de Bruijn and Hussain, 1984). H-GSP 8114 (and SZ; Raza et al.,
1984) have rodent faunas comparable to that of Potwar locality Y59Z. H-GSP 107,
8ZZ4, and 8ZZ7 all contain Antemus chinjiensis, known from 14.3 to 11.8 Ma in the
Potwar, and probably date to the older part of this range based on the presence of
Potwarmus primitivus in them as well (see Lindsay, 1987 and in press, for cricetid
ranges).
Figure 3 illustrates the detailed temporal durations, interpolated from strati-

graphic ranges, of Siwalik rodent taxa from 16.1 to 5.5 Ma. Small mammal sites and
their ages, from which figure 3 was constructed, are shown in table 1.
The Siwalik small mammal fauna is distinct from Europe throughout its history,
showing only sporadic and usually minor similarities to Europe relative to the total
rodent fauna known from a given time. Nevertheless, cricetids with European affin-
ities are basic to the development of the Siwalik muroid fauna. Murids are important
for correlations with Europe, but have not reached their full potential in this regard.
True gerbils have one Siwalik record at 7.5 Ma, whereas European records of its close
relative, Protatera, appear younger (see Aguilar et al., 1984, for example). The
pnrcupine Hystrix first occurs at about 7 Ma in Pakistan as an immigrant, as it does in
Europe. Sciurids and glirids may eventually prove useful for European correlations as
their systematic relationships become more fully known. Siwalik representatives of
other rodent families (e.g., Rhizomyidae, Ctenodactylidae, and Thryonomyidae) are
not directly relevant to European biochronology.
At the family level, cricetids dominate in diversity for two-thirds of the time
interval under discussion here (Lindsay, 1987). Between 16.1 and 1z. 7 Ma Siwalik
rodent faunas appear to resemble European faunas because they contain the cricetids
Megacricetodon and Democricetodon. This generic similarity is not particularly
informative for correlations because of the long temporal span in Europe of both
genera. However, it is significant in that these two genera are part of a post-Bugti
invasion that heralds the beginning of the Siwalik fauna per~· Other cricetid genera
are important for correlations with Africa and Asia Minor, notably Myocricetodon and
Dakkamys (Unay et al., 1985).
Murid rodents first occur at 14.3 Ma with Antemus. This is the oldest record of
murid rodents known anywhere in the world, based on a monophyletic definition of
murids as having two lingual cusps associated with the anterior two chevrons on the
upper first molar (Jacobs et al., in press). Similar cricetids are known from the same
time interval, including Potwarmus primitivus, a genus known also from the Miocene
of Thailand; however, no detailed stratophenetic transformation series of the evolu-
tion of murids from cricetids has been documented. The last record of Antemus is
11.8 Ma, based on a small sample with some similarities to Progonomys, including a
weak enterostyle-protocone (t4-t5) connection on one specimen.
Progonomys is first clearly present in the Siwaliks at locality Y634, a site nearly
10 km southwest of the locality with the youngest record of Antemus. Tracing beds
over this distance offers an imprecise correlation of ca. 11.6 Ma for Y634. The last
record of Antemus and the first record of Progonomys are close temporally. Based on
these Siwalik records, Progonomys is very likely to have evolved in South Asia, proba-
bly at approximately 11.8 Ma. Therefore, no European records are likely to be older
than 11.8 Ma. Regardless of its ancestry, the first occurrence of Progonomys in the
Siwaliks is notably older than that of Hipparion, rather than more-or-less synchronous
with it as in Europe.
In some features, early Siwalik Progonomys resembles Progonomys cathalai from

Europe and North Africa (Cheema et al., 1983). The last record of Siwalik
Progonomys is f.· debruijni at 7 Ma. The evolution of Mus from Progonomys is proba-
576
Ma
.' .
~
,_
·~
7 •"'
8 ~ I "'•
9
.
'-'
~·
~
"'"'
-~I
""
o= ~
10
"'
-l
11
12
13
I I~
14
"u
15
"'"'
c. :g .g
~
,_ ~
E
~
"
..
~
.X ~ ,_
~ ~
u
0
16 ~ ~
1+-RHIZOMYIDAE --+I MURIDAE +-----CRICETIDAE:-------+
Fig. 3. Potwar rodent biostratigraphy. Stratigraphic ranges of rodents from

the Potwar Plateau are plotted for sites readily tied to composite
paleomagnetic sections. Dots represent occurrences and vertical
lines show probable ranges. Question marks indicate specimens of
uncertain affinity and dashes indicate hypothetical ranges. Cricetid
data are derived from Lindsay (1987, in press); systematics of
Democricetodon and its allies are under study by E.H. Lindsay and,
for most lineages, only the first and last records are indicated by
dots. Data through 1987.
bly documented in the Siwaliks. The Siwalik earliest record of a taxon reasonably
included in the genus Mus is 5.5 Ma.
A change in the Siwalik murid fauna is recorded at about 8 Ma when

Paraethomys (see Brandy, 1981; originally reported as Parapodemus sp. by Jacobs,
1978) is recorded in Pakistan. Murids apparently dispersed between South Asia, North
Africa, and Europe at this time (Jacobs et al., in press).
577
~
24
20 +
Cll
' !_
+ ~
~ 16
u
UJ
0.. + ~
•
Cll 12
U-
II
0
t!
Ma a 9 io 11 12 13 14 15 16
Fig. 4. Histogram of rodent species at collecting horizons from 16.1 to 7 Ma.

Numbers tallied from figure Z for sites in table 1, and include both actual
and expected occurrences (see Barry et al., in press, for further discussion);
Cricetidae (unshaded), Muridae (diagonal shading), Rhizomyidae (black),
other rodents (horizontal shading). Larger samples (>150 specimens; arrows)
present the same pattern as all sites taken together.
Table 1. Potwar Plateau small mammal localities and ages from two time scales.
Localities are those used to construct figure z.
Age (Ma)
Mankinnen and Dalrymple Berggren et al.
Locality (1979) (1985)
DP-13 5.5 5.6

YGSP 437, 438 6.4 6.6
YGSP 369 6.5 6.7
YGSP 434 6.9 7.1
YGSP 547, 457 7.0 7.Z
YGSP Z4, 34, 17Z 7.Z 7.4
YGSP 387, 388 7.5 7.8
YGSP 367 7.8 8.1
YGSP 18Z, Z60, 410 8.0 8.4
YGSP Z70 8.Z 8.6
YGSP Z61, 330 8.5 8.9
YGSP 311 8.8 9.Z
YGSP 450 9.Z 9.6
YGSP Z59 9.4 9.8
YGSP 636 10.Z 10.6
YGSP 76 10.6 11.0
YGSP 504, 735 10.8 11.Z
YGSP 634 11.6 1Z.O
YGSP496 11.8 1Z.1
YGSP 668, 690, 691, 711, 7Z6 1Z.7 13.0
YGSP 41, 430, 640, 641 13.5 13.7
YGSP 665 13.7 13.9
YGSP 501 13.9 14.1
YGSP 589, 680 14.3 14.5
YGSP 709 14.7 14.9
YGSP 64Z 15.1 15.3
YGSP 591, 59Z 16.1 16.Z
578
APPARENT DIVERSlTY AND APPARENT TURNOVER
Figure 4 is a histogram of numbers of species recorded for all sites by age (table
1), including actual observations and range-through species (see Barry et al., in press,
for discussion of methods). Time intervals represented by larger samples (>150 speci-
mens) are indicated by arrows.
The total number of species is relatively stable among sites from 16.1 Ma until
13.7 Ma. Maximum diversity occurs at 13.5 Ma, after which diversity decreases,
including a sharp drop after 9.5 Ma. Extinction among cricetid& explains much of the
change in diversity between 16 and 7 Ma. Muridae are never particularly diverse.
Rhizomyids show some diversification by 8 Ma.
To examine turnover among rodents, numbers of first and last occurrences for
each locality in figure 4 were plotted (figure 5). A maximum of first appearances is
reached around 13.5 Ma, followed by a maximum in disappearances about one million
years later. Earlier and later periods of appearance and disappearance are less
marked.
Data become more directly comparable between sites when expressed as a

percentage of the total number of species known at well represented localities (figure
6). Middle Miocene maxima in appearances (13.5 Ma) and disappearances (1Z.7 Ma) are
less pronounced, but are still significant and diachronous (cf. figure 5). Major extinc-
tion, mainly among Cricetidae, occurs at 9.5 Ma. The late Miocene turnover is
evidently more severe than earlier events when considered relative to the total
number of rodent taxa. Families other than Cricetidae play a greater role.
The curves in figure 6 are smoother than those in figure 5 because only larger
samples (indicated by arrows in figure 4) are used. Data points representing small
samples are omitted as unreliable. For example, the low diversity (figure 4) of the 7.8
and 8.Z Ma sites is likely to be an artefact of small sample sizes. These sites also
show a lack of turnover events (figure 5) that may reflect small sample sizes, or
conversely, times of truly low turnover. By utilizing only higher quality data, sus-
12
•• 10
'ii
•... 8
•
0 6
•
~
..D
4
E
::J
z
2
0
7.0 Ma 8.0 9.0 10.0 11.0 12.0 13.0 14.0 15.0 16.0
0
•• 2
'ii
•... 4
•
0 6
•
~
..D
8
E
::J
z
10
12
Fig. 5. Numbers of species that are first occurrences (above) or last occur-
rences (below). See Barry et al. (in press) for a discussion of methods.
579
50
40
First Occurrences
30
%
20
10
7Ma 10 11 12 13 14 15 16
10
20
%
30
Last Occurrences
40
50
Fig. 6. First and last occurrences among larger samples (see figure 4) as per-
centages of the total microfauna. Peaks in first occurrences are at 13.5
and 8 to 7 Ma. Maxima in last occurrences are at 1Z. 7 Ma and after
9.5 Ma when extinction remains near 50% of the fauna.
tained high turnover in the late Miocene, particularly after 8 Ma, is seen to be the
fundamental pattern.
ABUNDANCE
The pattem of relative abundance changes illustrated by the biostratigraphic

record of Siwalik murids is informative when compared to that of cricetids. Currently
murids are the most diverse of living rodents, and they are generally considered to
have successfully competed with cricetids whenever such an encounter has occurred
(Misonne, 1969). While there are few cricetids in South Asia now, they appear to have
been quite diverse from about 16.1 to 9.5 Ma. Murids have never been diverse in the
portion of South Asia represented by the Siwaliks. Although they have not been par-
ticularly diverse, they have been abundant, judged simply by the relative percentages
of specimens recovered at screening localities (figure 7). Murids appear to achieve
consistent dominance in abundance over c:ticetids at about 11 Ma, even though the
earliest known record of murids is 14.3 Ma. Coexistence (and, inferentially, competi-
tion) between the two families occurred for over three million years before cricetids
relinquished dominance in abundance.
From 11 Ma onward, murid dominance of the rodent fauna is unchallenged

although there are some significant fluctuations as rhizomyids become temporarily
more abundant (their most abundant representation in the Siwaliks) between 8 and
7 Ma. The Brachyrhizomys (Rhizomyinae) lineage had achieved burrowing adaptations
by this time (Flynn, 198Z, 1986). Cricetids are not represented in known Siwalik local-
ities between 8.8 and 7.5 Ma even though the small mammal fauna is reasonably well
sampled in that interval. They temporarily reappear at 7.5 Ma.
While murid rodents are abundant in the Siwaliks, few samples have been
adequately described. Figure 8 shows histograms of the distribution of upper first
580
••
'Ill
-;:.
E
'I' rl' ''•---------·--------').-.

0
'I . . I
N
.1:
a: --·~
I ' , \ ,. '
,, _____ _
__ _
---
....
••
'Ill
••
..
'Ill
•
..
0
CJ
.
2
I
co ...
I • I
CD
A A A
Fig. 7. Muroid abundance in Siwalik assemblages (modified from Jacobs et al., in

press, fig. 9). Bars represent numbers of teeth as percentages of all teeth
recovered by locality (indicated by chevrons on left). Data through 1986. In
abundance, Muridae replace Cricetidae, while other taxa, including
rhizomyids, are generally uncommon.
molar length in undescribed Siwalik murid samples (abscissa) by interpolated age of

locality (ordinate). Described murid taxa are shown by horizontal lines representing
ranges of upper first molar length at localities YGSP 491, YGSP 18Z, and DP-13.
Upper molars of described taxa are illustrated in figure 9. Locality YGSP 491 has not
been precisely tied to a paleomagnetic section, but it is used here because it has the
largest and most adequately studied sample of Antemus (Jacobs et al., in press). It is
clearly as old or older than locality 496 based on associated fauna and lithostratig-
raphy. Samples used to construct figure 8 are listed in table z. Murid samples are
predominantly isolated teeth collected by sieving. Screen-wash localities are unevenly
distributed with clusters of sites at several time intervals.
The following features of figure 8 appear most striking: (1) size remains rela-
tively constant between 11.8 and 9.4 Ma; and (Z) increased size limits within Siwalik
murids becomes apparent by 8 Ma.
These features of figure 8 compared to figure 7 indicate that: (1) dominance of

murids over cricetids did not occur until after Progonomys evolved from Antemus;
(Z) striking increase in size of Siwalik Muridae occurs concomitantly with the
achievement of fossorial adaptations in rhizomyids and a brief reappearance of
cricetids. This is the time interval when the greatest percentage of taxonomic change
is taking place in the Siwalik fauna. The simultaneous affects on cricetid distribution,
rhizomyid burrowing habits, and murid body size between 8.5 and 7.5 Ma suggest that
abiotic factors are involved. A broad influence, such as climate, might be suspected.
The composite effect is reflected in the sustained high turnover rates of the late
Miocene seen in figure 6.
581
5.5
..... - -
••• • • ••• •
7.0 ~: •••• • .t.l.l .••.•
7.2
7.5
---I • •• .I. • I
8.0
• • • I •
-· -..
-;;-
6
<
Q)
0>0 8.8
• LL..
----- ......
9.2
9.4
• •
• 1•1 I I •
-·
-.
•
• •
• = I specimen
1.5 2.0 2.5

Length of First Upper Molar (mm)
Fig. 8. Murid first upper molar size through time. Horizontal bars indicate
observed ranges in published samples. Measurements for published
samples at 8 Ma (Progonomys debr1ni, Karnimata darwini, and
Parapodemus sp. ror Paraethomys fide randy, 1981]) and 5.5 Ma (Mus
auctor, Karnimata darwini, and Parapelomys robertsi) are from Jacobs
(1978). The horizontal bar at the bottom of the figure (13.5 Ma) repre-
sents the range of Antemus chinjiensis (from Jacobs et al., in press).
nlustrations of these taxa are shown in figure 9.
CONCLUSIONS
The Siwalik fauna represents a biogeographic region encompassing some or all of

South Asia. As such, its contributions to European biochronology are primarily in
constraining the timing of dispersal events. Large scale dispersal events are likely to
be related ultimately to abiotic factors such as climate and tectonics, in addition to
intrinsic biological attributes of organisms and the ecological relationships among
them. Such levels of complexity in biogeographic relationships require an increased
understanding of discrete geographic regions that relies fundamentally on an accurate
and consistent taxonomic data base, and on high resolution chronology.
582
Table z. Potwar Plateau sieving localities with murid
upper first molars (N) used to construct figure 8.
Locality Age (Ma) N
DP-13 5.5 49
YGSP 457 7.0 5
YGSP 547 7.0 5
YGSP Z4 7.Z 38
YGSP 34 7.Z 3
YGSP 387 7.5 z
YGSP 388 7.5 31
YGSP 367 7.8 10
YGSP 18Z 8.0 67
YGSP 311 8.8 15
YGSP 450 9.Z 6
YGSP Z59 9.4 Z5
YGSP 76 10.6 zz
YGSP 504 10.8 4
YGSP 634 11.6 4
YGSP 496 11.8 z
YGSP 491 3.Q.
Total 308
While detailed Siwalik chronology and in-depth systnmatic studies will continue
to enhance biochronologic correlations between South Asia and Europe, improvment in
precision based only on taxonomic similarity is likely to be mostly anecdotal and
unlikely to provide more than fine-tuning at this stage of studies. However, it is not
simply taxa that can be compared between Asia and Europe, but also the pattems of
evolution exhibited by them. This remains to be done, but it requires a refined and
independently calibrated chronology for Europe.
The establishment of a Siwalik fauna, distinct from that of the older Bugti
fauna, involved the dispersal of cricetids with European affinities into South Asia.
Cricetids diversified as they arrived in the area. Significantly, this diversification
gave rise to two still living monophyletic groups usually considered the families
Rhizomyidae and Muridae.
Siwalik cricetid diversity is greatly reduced after 9.5 Ma. Prior to that time,
cricetid composition is the major factor in the total Siwalik small mammal diversity.
After that time, turnover rates are sustained at high levels and involve all three
families.
Basic to the initial Siwalik cricetid radiation are the genera Democricetodon and
Megacricetodon. These two Miocene genera are wide-ranging, possibly with relatives
extending to North America. They, or their surrogates, have been implicated in the
ancestry of most living muroids (Jacobs and Lindsay, 1984). The most diverse family
of living mammals, the Muridae, has its roots in Siwalik rocks some 14.3 million years
old. Yet murids are not now, nor are they ever known to have been diverse in this
area of South Asia.
Antemus, the earliest known murid, apparently evolved in southern Asia between
18.3 and 14.3 Ma. Murids began to outcompete cricetid& with the evolution of
Progonomys, which occurred between 11.8 and 11.6 Main southern Asia. Then they
spread to other parts of the Old World where they underwent secondary radiations.
The most important dispersal event from South Asia to Europe among rodents is
that of Progonomys. Based on the Siwalik record, this event did not occur before
583
Fig. 9. Siwalik murid first upper molars (all specimens illus-
trated as if from right side; true position given
below): A, Antemus chinjiensis (Y7650), right; B,
Progonomys debruijni (Y7736), left; C, Karnimata
darwini (Y77ZO), right; D, Parapodemus sp. (Y7697) or
Paraethomys sp. (fide Brandy, 1971), left; E, Mus
auctor (DPZ10), left; F, Karnimata huxleyi (DPZ53),
left; G, Parapelomys robertsi (DPZ61), right. A from
locality YGSP 41 (13.5 Ma); B-D from locality YGSP
18Z (8.0 Ma); E-G from locality DP-13 (5.5 Ma). Bar
represents 1 mm. Lines indicate hypothetical trans-
formation series.
11.8 Ma. Judged from the Siwalik record, Progonomys, but perhaps not Antemus, may
have possessed the attributes of murids that facilitate their outcompeting cricetids.
Progonomys spread from South Asia to Europe and Africa where it was the base of
secondary murid radiations. The European murid radiation was significant, but it has
few living descendants (probably only Apodemus and Micromys). Modern murid divers-
ity, as a whole, is the result of repeated dispersal events to a number of separate
geographic areas at various times in the past, all of which postdate 11.8 Ma.
ACKNOWLEDGMENTS
We are grateful to the people of Pakistan, and for the opportunity to work in
their country. We very happily acknowledge the help and labors of our associates of
584
all nationalities who have been companions in field and laboratory. We would particu-
larly like to acknowledge Everett Lindsay and David Pilbeam. Lindsay guided two of
us through graduate school and over the years since then has continuously made avail-
able his help, comments, and unpublished data. Funding for our Siwalik work has come
from various NSF and Smithsonian grants. Todd Disotell and Charlotte Banasik helped
with illustrations. Pam Lunge drew the teeth in figure 9. Finally, we acknowledge
that the Siwalik record would not be understood so well if it were not for the chrono-
logie resolution based on magnetic polarity stratigraphy, due in very large part to the
dedicated efforts of our good friend and colleague Noye M. Johnson.
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l'Espagne) et le probleme de la migration messinienne. Paleobiologie
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Mirza, M.A., 1979. Magnetic polarity stratigraphy and vertebrate paleontology
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climatology, Palaeoecology, v. 27, p. 1-34.
geology and vertebrate fauna of the Miocene Manchar Formation, Sind, Potwar.
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Tauxe, L. and Opdyke, N.D., 1982. A time framework based on magnetostratigraphy
for the Siwalik sediments of the Khaur area, northern Pakistan. Palaeogeog-
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lating Miocene faunas from N. Africa, Turkey and Pakistan? 8th Congress of the
Regional Committee on Mediterranean Neogene Stratigraphy, Abstracts, p.
586-587.
Wessels, W., Bruijn, N. de, Hussain, S.T., and Leinders, J.J.M., 1982. Fossil rodents
from the Chinji Formation, Banda Daud Shah, Kohat, Pakistan. Proc. Kon. Ned.
Akad. Wetensch., ser. B, v. 85(3), p. 337-364.
586
THE AFRICAN DIMENSION IN EUROPEAN
EARLY MIOCENE MAMMAL FAUNAS
R. J. G. Savage
The University
Bristol BS8 lRJ, U.K.
INTRODUCTION
Ex Africa semper aliquid novi.
Out of Africa, always something new. So wrote Pliny almost ZOOO years ago. It
is still true today and it was also true Z4 million years ago at the beginning of Neogene
times. For it was out of Africa that there came the most distinctive markers in our
European faunas, the probscideans and the primates. Our task is to identify the
vanguard of the invasion and using those elements to trace them to reliably datable
deposits on the African continent. In East Africa the fossils are entombed in pyro-
clastic deposits. In consequence many sites can be radiometrically dated.
The principle is sound but its application is fraught with problems. In Africa the
earlier Neogene deposits are mostly in East Africa, a long way from the
Mediterranean and Europe; few, if any, of these can be dated back to the earliest
Miocene. The North African sites are not radiometrically datable, nor are the early
Neogene European sites. We have no alternative but to apply Occam's razor; the best
compromise remains the only solution.
When we re-examine the faunas from Kenya and Uganda we find not only a
wealth of information but also a sad lack of reliable data. Many of the collections
were made decades ago without adequate recording of field data; these early
discoveries are often the best specimens, but we may not know exactly where or from
which level the specimens came. The geology is often sedimentologically and struc-
turally complex, with the result that stratigraphical correlations are hazardous and
difficult. The radiometric sampling often gives conflicting dates, due in part to the
intense Pleistocene and post-Pleistocene weathering. There are many sites but there
is no one locality where a long succession of strata can be followed.
The object of this paper is to review the early Neogene African mammal faunas
insofar as they throw light on the problems of chronostratigraphically dating European
faunal assemblages. Included with these are the recently discovered faunas of Saudi
Arabia, for in early Miocene times this was part of the African continent. Also
included are the early Miocene faunas from Israel on account of their close proximity
to Africa and crucial position along possible migration corridors to Eurasia.

•
b\te"'
10 •
Fig. 1. Map showing the distribution of early Miocene

mammal localities in Africa and Asia Minor.
The scale of the map does not enable the many
sites in western Kenya and eastern Uganda to be
individually marked.
PALEOGEOGRAPHY
Before migration routes and distributions can be assessed, it is vital to know
something of the paleogeographical regimes. This data must come from independent
sources to avoid circular reasoning; this in practice means using sedimentological
data, floras, and marine faunas. The facies analyses will give us information on
environmental conditions and the marine faunas will establish the relative correlation
of sequences.
There is a well recognized pattern of steady regression of seas during Oligocene

times. One feature of this was the gradual closure of the Turgai Strait, allowing the
migration of Asian mammal families into Europe and producing the widely recognized
"Grande Coupure." The most notable feature of this great faunal turnover is the total
absence of African elements. All the sixteen families of mammals that make their
appearance in Europe in the early Oligocene can be traced to Asiatic origins. This
argues for the presence of a major Tethyan seaway stretching from the western
588
Mediterranean eastward to the Indian Ocean (Adams et al., 1983; Steininger et al.,
1985).
Bessedik and Sue (1983) argue on the basis of pollen for the persistence of
tropical vegetation with mangrove swamps and coral reefs in southern France in both
late Aquitanian and late Burdigalian times. During those periods island chains appear
to have arisen in the western Tethys, the Iberian peninsula began to rotate anticlock-
wise, and the Arabian platelet began to rotate on its collision course toward the
Turkish platelet. The foraminiferal evidence (Adams et al., 1983) indicates a break in
marine connections between the Mediterranean and Indian Ocean by mid Burdigalian
times. A product of this is the formation of the extensive marine evaporites in the
Lower Fars Formation of Iran (Steininger et al., 1985). The paleogeographic evidence
all points to the earliest Afro-Eurasian land links being established via Asia Minor in
early Burdigalian times at about Z1 Ma.
Our knowledge of the immediately pre-Neogene mammal faunas in Africa is

lamentably lacking. Our knowledge of Oligocene faunas is limited to those from the
early Oligocene of the Fayum in Egypt and a few scrappy sites across the Sahara. The
Fayum Oligocene fauna of land mammals from the Jebel el Quatrani Formation in
Egypt displays a highly endemic assemblage of tethytheres alongside primates,
creodonts, rodents, and anthracotheres. The latter groups must represent either
chance dispersals or earlier land links with Eurasia, possibly in late Eocene times in
the regressive phase following the Lutetian transgression.
EAST AFRICAN EARLY MIOCENE MAMMAL FAUNAS
The East African early Miocene mammal faunas will be considered first among
the African faunas. Although they are the most remote from Europe, the distance via
Ethiopia into Saudi Arabia and Asia is not so great. Also there is evidence that
savannah faunas extended northward in Miocene time right to the Mediterranean coast
with no Saharan desert barrier. The East African faunas are by far the best known and
the best dated of all African Miocene faunas. Although there are radiometric ages for
almost all the many sites in Kenya and Uganda, the spread of ages, allowing for error
margins, makes it impossible to place each site in precise chronologie order. Differ-
ences between two sites of apparently similar age but with differing faunal elements
may therefore be due to imprecision of dating resolution or to differences in the
ecological habitats of the sites.
A considerable step forward was made by Pickford (1981) in recogmzmg a

sequence of faunal assemblages and then age bracketing these. Table 1 summarizes
the faunal composition from Sets I and n which Pickford assigns to the early Miocene;
Pickford acknowledges that the differences between his Faunal Sets I and n may be
true time differences or may be due to biofacies differences. The only sites which fall
in Faunal Set I are Koru, Kiahera, Chamtwara, Legetet, Napak (lower levels), and
Songhor. The faunas from these sites are limited compared with the probably slightly
later and much richer Faunal Set II from Gumba, Moruorot, Chianda, Karungu, and
Hiwegi. Recently (Drake et al., 1988) have resampled some of the sites and give total
fusion K-Ar ages which are considerably younger than earlier assessments, and which
indicate that the sites are all much closer in age than had previously been believed.
The new ages for the Karungu, Kiahera, and Hiwegi sites indicate that all lie between
17.5 and 18.0 Ma; previously these sites were thought to range through about 7 Ma.
The new data thus indicates that the sites are late early Miocene (MN4). An impor-
tant extension to this work currently in hand is the reappraisal of the Set I sites.
There is, however, one site at Meswa Bridge which appears to considerably
predate the Set I faunas; the site has been dated at Z3.5 Ma and the European standard
stratigraphic table used in this volume places the base of the Miocene at Z4 Ma, a
redating of this site would be most important. The Meswa Bridge fauna comprises
Miorhynchocyon meswae, Proconsul major, Bathyergoides, Paranomalurus,
Paraphiomys, Megapedetes, Kelba quadeemae, Teratodon spekei, Hyaenaelurus sp.,
589
Table 1. Mammal fauna from sites in Faunal Sets I and IT of East Africa (Sources:
Drake et al., 1988; Pickford, 1981, 1986; Pickford et al., 1986; Schmidt-
Kittler, 1987).
Macroscelidea Myorhynchocyon clarki (Butler & Hopwood)

Myorhynchocyon meswae Butler
Myorhynchocyon rusingae (Butler)
Myohyrax oswaldi Andrews
Insectivora Galerix africanus Butler

Amphechinus rusingensis Butler
Gymnurechinus leakeyi Butler
Gymnurechinus camptolophus Butler
Protenrec tricuspis Butler & Hopwood
Parageogale aletris (Butler & Hopwood)
Erythrozootes champerpes Butler & Hopwood
Prochrysochloris miocenicus Butler & Hopwood
Chiroptera Propotto leakeyi Simpson

Taphozous incognita Butler & Hopwood
Hipposideros sp.
Chamtwaria pickfordi Butler
Primates Komba minor (Clark & Thomas)

Komba robustus (Clark & Thomas)
Progalago songhorensis Simpson
Progalago dorae Macinnes
Mioeuoticus sp. Leakey
Micropithecus clarki Fleagle & Simons
Dendropithecus macinnesi (Clark & Leakey)
Rangwapithecus g.ordoni (Andrews)
Nyanzapithecus vancouveringi (Andrews)
Limnopithecus legetet Hopwood
Proconsul africanus Hopwood
Proconsul major Clark & Leakey
Proconsul nyanzae Clark & Leakey
. Lagomorpha Kenyalagomys rusingae Macinnes

Kenyalagomys minor Macinnes
Rodentia Andrewsimys parvus Lavocat

Phiomys andrewsi Osborn
Paraphiomys pigotti Andrews
Paraphiomys stromeri (Hopwood)
Epiphiomys coryndoni Lavocat
Diamantomys luederitzi Stromer
Kenyamys mariae Lavocat
Simonimys genovefae Lavocat
Myophiomys arambourgi Lavocat
Elmerimys woodi Lavocat
Bathyergoides neotertiarius Stromer
Proheliophobius leakeyi Lavocat
Paranomalurus bishopi Lavocat
Paranomalurus soniae Lavocat
Paranomalurus walkeri Lavocat
Zenkerella wintoni Lavocat
Megapedetes pentadactylus Macinnes
Afrocricetodon songhori Lavocat
Protarsomys macinnesi Lavocat
Notocricetodon petteri Lavocat
Vulcanisciurus africanus Lavocat
590
Creodont a Kelba quadeemae Savage
Teratodon enigmae Savage
Teratodon spekei Savage
Anasinopa leakeyi Savage
Metapterodon kaiseri Stromer
Pterodon africanus Andrews
Pterodon nyanzae Savage
Leakitherium hiwegi Savage
Hyaenodon andrewsi Savage
Hyaenodon pilgrimi Savage
Megistotherium sp. Savage
Carnivora Hecubides euryodon Savage

Hecubides macrodon Savage
Hemicyon sp.
Luogale rusingensis Schmidt-Kittler
Kenyalutra songhorensis Schmidt-Kittler
Kichechia zamanae Savage
Legetetia nandii Schmidt-Kittler
Herpestides aequatorialis Schmidt-Kittler
Leptoplesictis rangwai Schmidt-Kittler
Leptoplesictis mbtensis Schmidt-Kittler
Mioprionodon pickfordi Schmidt-Kittler
Stenplesictis muhoronii Schmidt-Kittler
Afrosmilus africanus (Andrews)
Afrosmilus trukanae Schmidt-Kittler
Tubulidentata Myorycteropus africanus Macinnes

Orycteropus minutus Pickford
Proboscidea Prodeinotherium hobleyi (Andrews)

Eozygodon morotoensis (Pickford & Tassy)
Archaeobelodon aff. filholi Tassy
Gomphotherium sp.
Platybelodon sp.
Hyracoidea Pachyhyrax cham ioni (Arambourg)

Prohyrax bateae hitworth)
Perissodactyla Chalicotherium rusingense Butler

Aceratherium acutirostratum (Deraniyagala)
Brachypotherium heinzelini Hooijer
Dicerorhinus leakeyi Hooijer
Artiodactyla Hyoboops africanus (Andrews)

Brachyodus aequatorialis Macinnes
Xenochoerus africanus (Stromer)
Libycochoerus jeanelli (Arambourg)
Kenyasus rusin ensis Pickford
Kenyasus kijivium Wilkinson)
Dorcatherium chappuisi Arambourg
Dorcatherium pigotti Whitworth
Dorcatherium parvum Whitworth
Dorcatherium songorhensis Whitworth
Canthumeryx sirtensis Hamilton
Propalaeoryx nyanzae Whitworth
Walangania africanus (Whitworth)
Gelocus whitworthi Hamilton
591
Orycteropus minutus, Eozygodon morotensis, Anthracotheriid, and Dorcatherium
songhorensis (Pickford, 1986).
The most significant records are the presence in these East African faunas of
proboscideans; Eozygodon morotoensis at Meswa, Archaeobelodon sp. at Songhor and
Legetet, Prodeinotherium hobleyi at Koru (Tassy, 1986). These primitive probo-
scideans represent the earliest occurrence of the group on the continent. The Faunal
Sets I and II share with the Agenian sites of Europe (MN1 and MNZ) a number of
mammalian families; these include Amphicyonidae, Viverridae, Ochotonidae,
Theridomyidae, Sciuridae, Cricetidae, Rhinocerotidae, Anthracotheriidae, and
Suidae. The major problem at present, however, is to resolve the time differences
between Set I and Set IT. Faunal Set IT has also many affinities with the early
Orleanian (MN3) though the radiometric dating suggests MN4.
MIOCENE MAMMAL FAUNAS FROM GEBEL ZELTEN, LIBYA
North of the Sahara there are no known early Miocene sites in Morocco, Algeria,
or Tunisia, though a few localities have yielded scraps of proboscideans which indicate
a Miocene age in the broad sense. From Libya and Egypt, however, there are good
faunas which can be placed firmly in the early Miocene, though their exact correlation
is still subject to review. The richness of these Miocene faunas indicates a tropical
regime with open shrubland faunas; there is no evidence of a desert barrier between
central Africa and the Tethyan shores.
In Libya the major sites are along the southern escarpment of the Gebel Zelten.
Selley (1966) recognized in the Marada Formation a series of clastic facies each with a
characteristic lithology, environment, and fauna; he envisaged the sequence as
comprising offshore bars with lagoons behind, into which issued rivers depositing
fluvial sediments. The river flowed north through what is now Gebel Zelten, with
distributaries on the north side of the escarpment. The reptile and mammal faunas
mostly come from coarse bedded sands and shales of the fluviatile members.
The Marada Formation at Marada rests on the Gier Bu Hasciso Formation, a

marine sequence with Nummulties intermedius fitcheli, which is characteristic of
Rupelian age in North African sequences. The Dor Zaggut sections of the Marada
Formation contain shelly limestones with Borelis melo (N9-N1Z) and Borelis melo
curdica (N8-N9). Perhaps less reliable but nevertheless consistent evidence comes
from the shelly fauna; marine bands can be traced south into Gebel Zelten within the
Marada Formation and in these the oyster reefs of Crassostrea gryphoides are common
and widespread. The range of this oyster is regarded as characteristic of a marine
transgression in N6 (V1m Couvering and Berggren, 1977). Most of the mammals found
on the south side of the Gebel Zelten occur in beds below the marine transgression
with oysters.
The Moghara Formation of Egypt is the eastward equivalent of the Marada

Formation and has a fauna which is closely comparable with that of Gebel Zelten (see
below and Said, 1962).
The mammal fauna from Gebel Zelten (table Z) contains a rich mixture of stocks
indicating communication with central Africa and across Tethys with Eurasia. There
are proboscideans, creodonts, carnivores, rhinoceroses, suids, and bovids which are
shared at generic level with East Africa. Fejfar (pers. comm.) has recently found
rodents at Gebel Zelten wh~h at generic level suggest comparison with middle and
late Miocene levels in the Siwaliks.
Savage and Hamilton (1973) concluded that the fauna was probably !~lightly
earlier than the mid Burdigalian faunas of France. Pickford (in press) has compared
the Gebel Zelten fa,una with faunas from sites in East Africa, Tunisia (Beni Mellal and
Bou Hanifia), and Egypt (Moghara and Siwa). He concludes the strongest correlation is
with Maboko in Kenya, a fauna he places near the base of his Faunal Set IIIb and
592
Table z. Mammal fauna from sites in the Gebel Zelten, Libya (Sources: Fejfar, pers.
comm.; Pickford, 1987; Savage and Hamilton, 1973; Savage, unpublished).
Primates Prohylobates simonsi Delson
Lagomorpha Ochotonidae indet.
Rodentia Diatomys sp.

cf. Potwarmus sp.
Democricetodon spp.
Africanomys sp.
cf. Dakkomys sp.
Zapodidae indet.
Creodont a Megistotherium osteothlastes Savage

Anasinopa sp.
Carnivora Afrocyon burolleti Arambourg

Syrtosmilus syrtensis Ginsburg
Proboscidea Prodeinotherium hobleyi (Andrews)

Gomphotherium angustidens (Cuvier)
Gomphotherium pygmaeum Arambourg
Hyracoidea Saghatheriine cf. Prohyrax
Perissodactyla Brachypotherium snowi (Fourtau)

Aceratherium campbelli Hamilton
Artiodactyla Brachyodus sp. nov.

Sivameryx africanus (Andrews)
Xenochoerus africanus Stromer
Libycochoerus massai (Arambourg)
Libycochoerus khinzikebirus (Wilkinson)
Listriodon akatidogus Wilkinson
Canthumeryx sirtensis Hamilton
Prolibytherium magnieri Arambourg
?Palaeomeryx sp.
Dorcatherium libiensis Hamilton
Eotragus sp.
Protragocerus sp.
equates it with MN4a. When we compare the faunas of Gebel Zelten with those of
East Africa, we find about the same degree of similarity with Faunal Set 1/II and
Faunal Set illb (characterized by the Maboko fauna). The Zelten fauna has nine genera
and five species in common with Set 1/II fauna and eight genera and three species in
common with Set IIIb. However, the three species which the Zelten fauna shares with
Maboko (i.e., Prodeinotherium hobleyi, Xenochoerus africanus, and Canthumeryx
sirtensis are also present in the earlier East African Set 1/II. In addition, three genera
are known from all three levels (i.e., Anasinopa, Brachypotherium, and Dorca-
therium). We are thus left only with a couple of suids, Libycochoerus khinzikebirus
and Listriodon akatidogus which appear to be shared by the Zelten and Maboko
faunas. However, the taxonomic identity of African suids (and anthracotheres) is very
shaky; every paper published in recent years has produced yet another horrendous
change of names and the synonomy now surpasses that of any other mammalian stock
on the continent. A careful and responsible taxonomic revision for both suids and
anthracotheriids is sorely needed.
Thus we have as yet no firm resolution of age for the Gebel Zelten fauna. A
broadly Orleanian age is agreed. Comparison with East Africa does not resolve the
593
problem further. Analysis of the new rodent fauna could give us a breakthrough.
Taking all the evidence -- lithological, marine faunas, and mammals - the best
estimate is late MN3b.
MIOCENE MAMMAL FAUNAS OF EGYPT
Hamilton (1973a,b) has given the most up-to-date revtston of the faunal lists
(table 3) for the early Miocene mammals known from three sites in Egypt. Their yield
is small but they provide a useful link geographically with the faunas of Israel and
Saudi Arabia. Geologically, the sites occur in fluviatile beds which appear to be the
eastern continuation of the Marada Formation of Libya. The faunas are all very
similar; all the genera known from Moghara are present at Zelten and in several cases
the species are probably identical. Siwa and Wadi Faregh faunas are impoverished
versions of the Moghara fauna with no new species to add to the list. There are thus
good grounds for equating these sites with Gebel Zelten in age.
Table 3. Mammal fauna from Miocene sites in Egypt (Sources: Hamilton, 1973a,b).
Wadi
Moghara Siwa Faregh
Primates Prohylobates tandyi Fourtau X
Creodonta Hyainailouros fourtaui von Koenigswald X
Proboscidea Gomphotherium angustidens (Cuvier) X X

Gomphotherium spenceri (Fourtau) X X
Perissodactyla Brachypotherium snowi (Fourtau) X X

Aceratherium campbelli Hamilton X
Artiodactyla Brachyodus africanus Andrews X X X

Brachyodus depereti (Fourtau) X X
Brachyodus sp. X
Sivameryx moneyi (Fourtau) X
MIOCENE MAMMAL FAUNA OF ISRAEL
Tchernov et al. (1987) have described the mammal fauna from the Rotem and
Yeroham basins in the Negev of Israel (table 4). These two basins contain fluviatile
coarse grained sands. In the Yeroham basin these are overlain by transgressive marine
beds containing oyster reefs with Crassostrea gingensis; this transgressive phase is
regarded by the authors as an early Miocene event (N4-N7).
The fauna is small but has its strongest affinities with the early Miocene sites of'
East Africa. All the genera, save only one rodent (Metasayimys), are known in East
African sites of early Miocene age (see table 1); in many cases there is even specific
identity. The genus Metasayimys is recorded from Saudi Arabia (see below). The
presence of proboscideans, creodonts, carnivores, and bovids with close East African
affinities is surely witness to good land connections between the two areas and prob-
ably relatively similar environmental conditions. There is no evidence to suggest that
the age of the Negev fauna is different from that of the early Miocene faunas in East
Africa, but the imprecision of dating on the East African faunas leaves a wide error
margin. There is also a strong affinity with the Gebel Zelten fauna, especially among
the artiodactyls. Tchernov et al. (1987) while noting the similarities of the Negev
fauna to those from Gebel Zelten and Moghara conclude that, on the basis of all the
geological and faunal evidence, the Negev mammal fauna is probably closest in age to
those of MN3a in Europe.
594
Table 4. Mammal fauna from the Miocene of Israel (Source: Tchernov et al., 1987).
lnsectivora Erinaceidae, gen. indet.
Primates Dryopithecinae, gen. indet.
Lagomorpha Kenyalagomys sp.
Rodentia Megapedetes cf. pentadactylus Macinnes

Metasayimys sp.
Cricetidae, gen. indet.
?Bathyergidae, gen. indet.
Creodont a Anasinopa haasi Tchernov
Carnivora Viverrinae, gen. indet.
Proboscidea Prodeinotherium sp.

Gomphotherium sp.
Perissodactyla cf. Dicerorhinus sp.
Artiodactyla Listriodontinae, gen. indet.

cf. Canthumeryx sirtensis Hamilton
Dorcatherium cf. pigotti Whitworth
Dorcatherium cf. cha uisi Arambourg
Eotragus cf. sansaniensis Lartet)
Gazella negevensis Tchernov
cf. Walangania
MIOCENE MAMMAL FAUNAS OF SAUDI ARABIA
Mammal faunas are found in Saudi Arabia at three stratigraphic levels; the
Hadrukh Formation, Dam Formation, and Hofuf Formation.
The Hadrukh Formation is the oldest of the three mammaliferous levels; it

comprises continental deposits which unconformably overlie marine Eocene
sequences. In the absence of planktonic foraminifera, the precise age of the Hadrukh
beds is uncertain and estimates vary from Chattian to Burdigalian (Whybrow, 1987).
From the locality of Jabal Midra ash-Shamali, a small vertebrate fauna has been
identified. The mammals are very rare, comprising only a bovid (cf. Oioceros sp.) and
two rodents; these are the zapodid Arabosminthus quadratus and t,he cricetid
Shamalina tuberculata (Daams, 1982). These do not help date the deposit; they merely
indicate a broadly middle Miocene age.
The marine carbonates of the Dam Formation have a fossiliferous continental

equivalent at Ad Dabtiyah and As Sarar. The fossils occur in fluvial sandy carbonate
muds with discontinuous channel conglomerates. Freshwater fish, turtles, and croco-
diles are found along with the terrestrial mammals.
Whybrow (1981) and Thomas et al. (1982) have given the most recent account of
the mammal faunas from the Dam Formation sites in Saudi Arabia (table 5); the table
combines the faunal evidence collected by the British team under Whybrow working at
Ad Dabtiyah and the French team working under Thomas at As Sarar. The zoogeo-
graphic affinities of the fauna are not clear; many share similarities with East Africa
and with Pakistan, suggesting communication routes were in existence between the
continents. None of the mammals enable a precise age determination to be given; the
closest comparison in Kenya is with Maboko. The overall concensus is that the fauna
is mid Orleanian, about MN4a.
595
Table 5. Mammal fauna from two Miocene sites in the Dam Formation, Saudi Arabia
(Sources: Thomas et al., 198Z; Whybrow, 1987; Whybrow et al., 198Z).
Ad As
Dabtiyah Sarar
Primates Heliopithecus leakeyi Andrews & Martin X

gen. & sp. indet. X
Rodentia Metasayimys cf. intermedius Sen & Thomas X

Megapedetes cf. pentadactylus Macinnes X
cf. Protalactaga X
Paraphiomys sp. X
Carnivora Viverra sp. X

cf. Martes X
Mionictis sp. X
Pseudaelurus tournauensis (Hoernes) X
Amphicyon sp. X
Proboscidea cf. Deinotherium X

Gomphotherium cooperi (Osborn) X
Gomphotherium sp. X
Hyracoidea Pachyhyrax aff. championi (Arambourg) X
Perissodactyla Aceratherium sp. X

Brachypotherium sp. X
Dicerorhinus sp. X X
Artiodactyla Listriodon sp. X X

?Kenyasus X
Canthumeryx sp. X
Dorcatherium sp. X X
Eotragus sp. X
The Hofuf Formation is the youngest of the three levels with mammal faunas.
Fossils have been collected from a small gravelly sand channel at AI Jadidah. In
addition to fish, chelonians, and crocodiles, the following mammals are recorded (Sen
and Thomas, 1979; Thomas et al., 1978; Thomas, 1983):
Rodentia Sciuridae Atlantoxerus sp.

Ctenodactylidae Metasayimys intermedius Sen & Thomas
Carnivora Hyaenidae Percrocuta sp.
Proboscidea Gomphotheriidae Gomphotherium angustidens (Cuvier)
Perissodactyla Rhinocerotidae Dicerorhinus cf. primaevus Arambourg
Artiodactyla Suidae cf. Lopholistriodon

Giraffidae Palaeotragus sp.
Bovidae Pachytragus ligabuei Thomas
Caprotragoides aff. potwaricus (Pilgrim)
Protragocerus sp.
cf. Homoiodorcas?
Thomas (1983) describes the bovids in detail and interprets them as evidence of
immigration influences from Asia Minor (e.g., Pachytragus) and from Africa (e.g.,
Caprotragoides and Homoiodorcas). The closest similarities to this fauna are to be
596
found in those of Fort Ternan in Kenya (14 Ma) and Beni Mellal in Morocco which he
places in MN7. The AI Jadidah fauna is pre-Hipparion and on the basis of all the
available evidence Thomas therefore places it at the MN6/MN7 boundary.
CONCLUSIONS
This review brings up to date the information on early Miocene mammal faunas
in Africa and adjacent parts of Asia Minor. New radiometric dating analyses of the
Kenyan sites give results which are helping to place the sites in a more precise strati-
graphic succession. Apart from the fauna at Meswa Bridge which is possibly around
24 Ma, the redated early sites all group around 18 Ma. If the European correlation of
dating with mammal levels is correct, then the early Kenyan faunas belong to MN4,
with the exception of Meswa which falls in MN2. However, the sites chosen for
redating are essentially those of Faunal Set II and as yet we have no update on the Set
I sites, in particular for Songhor. It could be expected that these, if the faunal differ-
ences are true time differences, will yield considerably older dates; the affinities of
the Set I fauna are more with the early Orleanian zone of MN3 than with MN4.
The Libyan fauna from Gebel Zelten has similarities not only with the early
Kenyan faunas of Set II, but also with the younger fauna from Maboko. The marine
shelly fauna suggests a date close to the MN3b/MN4 boundary and this is not in
conflict with the mammal evidence. The Egyptian sites of Moghara and Siwa must be
similar in age to Zelten.
The Israeli mammal sites in the Negev have much in common with Zelten, both
in faunal affinities and in geological relationships. It is difficult to make a case for a
different age from Zelten.
From Saudi Arabia the Dam Formation sites fall close to those of Zelten and
Maboko, with the Hadrukh fauna older. The Hofuf fauna is much younger, though still
pre-Hipparion, and is comparable to Fort Ternan in Kenya (14 Ma), which falls on the
MN6/MN7 boundary.
The difficulties we experience in attempting these correlations are almost

entirely due to our ignorance. As we acquire a larger database, so more surely will we
be able to place the faunas in a truly chronological sequence.
Table 6. Correlation of early Miocene mammal faunal levels in Africa and Asia
Minor.
MN Zones East Africa Libya ~ Israel Saudi Arabia
7
Fort Ternan Hofuf Fm.
6
4b
Maboko
4a Dam Fm.
Zelten Moghara-Siwa Negev
3b Faunal Set II
?Hadrukh Fm.
3a Faunal Set I
2 Meswa
597
ACKNOWLEDGMENTS
The author is grateful to the organizers of the NATO workshop for the invitation
to attend the meeting at Schloss Reisensberg and to Drs. Fahlbusch and Lindsay for
the opportunity to make a contribution to the proceedings on the African faunas.
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Event. Utrecht Micropal. Bull., v. 30, p. 273-298.
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Colloquium, Montpellier, Avril1983, p. 33-37.
Daams, R., 1982. Rodents from the early Miocene of eastern Saudi Arabia, in
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Pickford, M.H.L., Senut, B., Hadoto, D., Musisi, J ., and Kariira, C., 1986. Nouvelles
decouvertes dans le Miocene inferieur de Napak, Ouganda Oriental. C.R. Acad.
S. Paris, 302, serie II, p. 47-52.
Said, R., 1962. The Geology of Egypt. Elsevier, Amsterdam.
Savage, R.J.G. and Hamilton, W.R., 1973. Introduction to the Miocene mammal
faunas of Gebel Zelten, Libya. Bull. Brit. Mus. Nat. Hist. (Geol.), v. 22, p.
515-527.
Schmidt-Kittler, N., 1987. The Carnivora (Fissipedia) from the lower Miocene of East
Africa. Palaeontographica Abt. A., v. 197, p. 85-126.
Selley, R.C., 1966. The Miocene rocks of Marada and the Jebel Zelten: A study of
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Sen, S. and Thomas, H., 1979. Decouverte de Rongeurs dans le Miocene moyen de la
Formation Hofuf (Province du Rasa, Arabie saoudite). C.R. somm. Soc. geol.
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Tassy, P., 1986. Nouveaux Elephantoidea (Mammalia) dans le Miocene du Kenya.
C.N.R.S. Cahiers de Palaeontologie.
Tchernov, E., Ginsburg, L., Tassy, P., and Goldsmith, N.F., 1987. Miocene mammals
of the Negev (Israel). J. Vert. Paleont., v. 7, p. 284-310.
Thomas, H., 1983. Les Bovidae (Artiodactyla, Mammalia) du Miocene moyen de la
Formation Hofuf (Province du Rasa, Arabie saoudite). Palaeovertebrata,
Montpellier, v. 13, p. 157-206.
Thomas, H., Taquet, P., Ligabue, G., and Del'Agnola, C., 1978. Decouverte d'un
gisement de Vertebres dans les depots continentaux du Miocene moyen du Rasa
(Arabie saoudite). C.R. somm. Soc. geol. Fr., 1978 fasc. 2, p. 69-72.
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Thomas, H., Sen, S., Khan, M., Battail, B., and Ligabue, G., 198Z. The lower Miocene
fauna of Al-Sarrar (Eastern Province, Saudi Arabia). ATLAL, Jl. Saudi Arab.
Archaeol. Jeddah, v. 5, p. 109-136.
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Neogene time scale, in Kaufman, E.G. and Hazel, J.E. (eds.), "Concepts and
Methods of Biostratigraphy." Dowden, Hutchison and Ross, Inc., Stroudsbourg.
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599
DEVELOPMENT AND APPUCATION OF LAND MAMMAL AGES IN
NORTH AMERICA AND EUROPE, A COMPARISON
Everett H. Lindsay
Tucson, Arizona 85721, U.S.A.
Richard H. Tedford
Department of Vertebrate Paleontology

American Museum of Natural History
New York, New York 10024, U.S.A.
INTRODUCTION
During the last century the geologic time scale was developed from a sequential
array of fossiliferous marine deposits, primarily in Europe. Prior to the last century
the concept of geologic time was addressed by numerous natural scientists, including
Steno, Buffon, Hutton, and Smith. With subsequent refinements, this time scale is now
widely accepted as the chronologie framework for all geologic and biologic events in
earth history. However, with rare exceptions (e.g., the Paris Basin) the application of
this framework to continental sediments has been difficult because continental
deposits are usually less extensive, superposition is more difficult to demonstrate, and
interdigitation of marine and nonmarine rocks are rare. Because of these factors,
vertebrate paleontologists have made few contributions to the development of the
geologic time scale, even though their contributions to the concept of organic
evolution has been significant.
Vertebrate paleontologists have developed their own chronologie system, based

primarily on the evolution of terrestrial mammals. This ordinal system has been tied
to the geologic time scale partly by reference to marine biostratigraphic units where
interdigitating marine deposits are available, and partly using radiometric and
paleomagnetic data. This terrestrial chronologie system, known as regional (or
continental) land mammal ages, has proven reliable and divisible to the same degree as
the marine stratigraphic sequence that gave rise to the geologic time scale. Perhaps
it is anomalous that land mammal ages, and the principles on which they are founded,
have never been identified or included within any of the stratigraphic codes used by
stratigraphers and geochronologists. The terms "continental," "mammal" and "terres-
trial" are not indexed in the International Stratigraphic Guide (Hedberg et al., 1975).
It is even more anomalous that vertebrate paleontologists have not argued for
recognition, within existing stratigraphic codes, for concepts and terminology applica-
ble to land mammal ages. Land mammal ages are not an offshoot or "secondary"
system of less refined paleontologic-stratigraphic methods compared to those applied
in marine sequences; nor should they be regarded as the unwanted offspring of marine
stratigraphers.

Geographic Limits of Land Mammal Ages
Land mammal ages are widely recognized as a reliable chronologie system for
terrestrial rock sequences deposited during the Cenozoic. However, it should be
emphasized that each continent was, at different times, isolated from the others
during most of the Cenozoic; therefore, land mammal ages, which reflect mammalian
evolution within separate zoogeographic regions (or continents), are different within
each continent or biogeographic region. Mammalian history is punctuated by the
introduction of immigrants from other regions. It is commonly the infrequent
introduction of immigrants, e.g. dispersal events, that permit recognition of discrete
segments of faunal history within each region, and that also aid in the correlation of
land mammal ages in separate regions. Thus, mammals on each continent or region
have their own distinctive'history, and there is no~ priori reason why the sequence of
events on one continent should be coincident with the sequence of events on other
continents or with divisions of the geologic time scale. However, the geologic time
scale, as developed and applied in marine rock sequences (and calibrated by
radiometric and paleomagnetic methods) provides the standard chronologie framework
for correlating all of Earth history, including mammals.
This report is an overview of Cenozoic mammalian biochronology, emphasizing

its current status in North America and Europe, where mammalian biochronology is
framed in land mammal ages, stages, or mammal zones, and where the known record
of mammal evolution is more complete. A second purpose is to compare the develop-
ment of land mammal ages in North America and Europe, to identify steps leading
toward the proposal of land mammal ages, and to clarify remaining problems that
must be addressed before a global chronologie framework for mammal evolution can
be realized.
NORTH AMERICAN LAND MAMMAL AGES
Land mammal ages were conceived in North America and have been tested
longer and more thoroughly there than elsewhere. The concept of land mammal ages
comes from the "life zones" of Osborn and Matthew (1909). These paleontologists
described "Cenozoic mammal horizons of western North America" almost 80 years
ago. They clearly stated in their opening sentence, "The main purpose of this paper is
faunistic rather than geologic" (Osborn and Matthew, 1909, p. 7). A central theme of
their study was the comparison of "two natural divisions" (e.g., the Mountain Region
and the Plains Region) within the Cenozoic deposits of North America. They
considered principles (pages 19-3 Z), then proceeded to describe seven faunal phases
(comprised of 19 mammal life zones) as temporally discrete units of "a nearly com-
plete and unbroken history of the Tertiary" in North America, based on representative
fossil mammals, from restricted continental deposits in well known stratigraphic
sequences.
Some of the fossil deposits are superposed but most are not, and direct
superposition of fossil sites is rare. Figure 2 (from Osborn and Matthew, 1909) shows
superposition of some Eocene faunas in separate basins of Wyoming. These sites were
instrumental in developing the stratigraphic-faunal sequence of life zones by Osborn
and Matthew. Geographic distribution of fossil deposits in North America is also
restricted, with the same deposit rarely occurring in more than one state. However,
the concensus at that time called all discontinuous early Eocene terrestrial redbed
sequences in North America the Wasatch Formation as though this deposit represented
a nearly continuous sedimentary "blanket" over widespread areas and could be utilized
as a stratigraphic marker, enclosing distinctive fossils (e.g., Hyracotherium,
Coryphodon, Pelycodus, etc.) in much the same way as a widespread marine deposit
would be utilized.
The study by Osborn and Matthew resulted in a sequential ordering of relatively

isolated continental sediments, correlated by the distinctive fossils contained in the
sediments. Divisions of this framework were called "life zones," named for a char-
602
North American Land European European
Mammal Ages Mammal Ages Mammal Stages
Geologic Time
(Woadburne, 1987) Scale (fahlbusch, 1976) (from Van de Weerd, 1976;
Daams &: freudental, 1988)
Rancholabrean Ma Pleistocene Ma
Irvingtonian 0.5 ~ Biharian
1.5 1.6
Blancan Vailanyian
4.5 Pliocene Ruscinian
5.0
-- Hemphillian 1
8.0 Turolian Turolian
Clarendonlan Vallesian Vallesian
11.5
Barstovian Miocene Astaracian

16.5 Aragonian
Hemingfordian
20.0 Orlean ian
Rambllan
24.0 Agenian
Arikareean
29.0 Arvernian
Whitneyan
31.0
Orellan Oligocene
-· 32.0 Suevian
Chadronian
38.0 Headonian
39.0
Duchesnean .42.0
Unitan Rhenanian
48.5
Eocene
~
Bridgerian 51.0
Wasatchian Neustrian
~
57.5
Clarkforkian ~
59.5 58.5
Tiffanian I
63.0 Paleocene
Torrejonian Paleocene
I 5.0
Puercan 66.5 66.5
(!)
0 Fig. 1. North American land mammal ages, European mammal ages, and European mammal stages.
w
Coryp}wJDn Tn;,~s:~oM!pu"::~n,1,
6.t>IV "·~~,.,.,.,.,.,
..
... •~r•d;.nrt
., ·
•
· ~~
l~ticqu
..
~zone
hlfl:od"s
Oxy«.,.
AN>eod<N>
£ohippu• (~ryllbuml.nt)
Sy;stemot/()n
CtNyphodon c~,,!Jund4ttt)
Sino~
Th~sU#
Hyop~rxlu•
S.I"CCO~mur
E3thonYJI
h/Hk:top$.
f'llf111'fly3
...
PI.,UIIM
il'tWf't•..,.~tu
()/'M~~nt:.,.
Fig. Z. Composite stratigraphic sections of the Wasatch Formation, from the

Bighorn Basin in northern Wyoming (left side) and the Bridger Basin in
southern Wyoming (right side). (Figures 3 and 4 from Osborn and
Matthew, 1909.)
acteristic and well represented mammal genus from that life zone. Figure 3 (Figure
10 in Osborn and Matthew, 1909) shows how stratigraphic sequences with representa-
tive fossils from more widespread areas of North America were assembled by Osborn
and Matthew to develop the younger life zones. Osborn and Matthew considered the
chief remaining temporal gap within the North American sequence occurs in the
Pliocene, or between zone 17 (the Glyptotherium zone) and zone 18 (the Elephas
imperator zone). We now recognize these zones as the late Pliocene Blancan and early
Pleistocene Irvingtonian land mammal ages. It is interesting that this faunistic
framework was correlated with marine stages of Europe rather than with marine
deposits of North America. These correlations were appropriate because European
continental deposits were also placed within a framework of European marine stages,
and marine-nonmarine tie-ins were both rare and poorly described in North America at
that time. Figure 4 (from Tedford, 1970) illustrates the development of the life zone
concept by Osborn and Matthew, from the time of Hayden (1869). Note that the
faunal zones of Matthew (19Z4) do not permit as much chronologie resolution as the
life zones of Osborn and Matthew (1909) as they are based on a single lineage, the
horse family.
Stratigraphic Codes and LaDd Mammal Ages
The development of stratigraphic codes went hand-in-hand with the development

of continental life zones, as proposed by Osborn and Matthe w. The first stratigraphic
code was conceived in discussions at the Seventh International Geological Congress,
held at Paris in 1897. Pressure to develop a more comprehensive and explicit strati-
graphic code continued, and in 1933 the first North American stratigraphic code was
604
Fig. 3. Relative placement of Oligocene-Pleistocene stratigraphic sequences of
western North America, to order the younger North American life
zones. (Figure 10 from Osborn and Matthew, 1909.)
Fig. 4. Comparative construction of mammalian faunal units in North

American terrestrial deposits, from Hayden (1869) to Matthew (19Z4).
(Figure 1 in Tedford, 1970.)
published. Vertebrate paleontologists in North America were eager to keep their

continental chronologie framework consistent with current practice and philosophy.
At the 1937 meeting of the Vertebrate Section of the Paleontological Society, the
Chairman, Walter Granger, appointed a committee of seven persons to canvas the
utility of a provincial North American Tertiary time scale, and if seen fit, to formally
propose and publish such a provincial time scale. A committee report was presented
at the 1938 annual meeting; it was agreed to extend the study for wider distribution
and critique. A 16-page report was distributed prior to the 1939 annual meeting; it
605
was debated and revised before final acceptance at that meeting, and published in
1941 in the Bulletin of the Geological Society of America (Wood et al., 1941).
Wood Committee Report
The 1941 publication, defining formal North American Provincial Land Mammal
Ages, is commonly referred to as the Wood Committee Report, in honor of the chair-
man of that committee, Horace E. Wood n. The term "provincial" is now commonly
dropped, as it was really intended to apply to all of North America which is more
appropriately identified as a biogeographic region, and redundant in the title. The
Wood Committee Report was written to conform with the 1933 North American
Stratigraphic Code, but it was completely ignored in the next edition of the North
American Stratigraphic Code (in 1961), which included numerous improvements over
the earlier 1933 Stratigraphic Code. Apparently, paleontologists active in drafting the
stratigraphic codes have always considered that continental rock sequences can and
should conform to the concepts and terminology applicable to marine rock sequences.
In other words, land mammal ages could eventually be incorporated into chronostrati-
graphic units, more or less equivalent in concept to marine stages. However, neither
vertebrate paleontologists nor marine stratigraphers would be likely either then or
now to accept the equivalence of land mammal ages and marine stages.
The Wood Committee Report named 17 land mammal ages, plus naming an older
Lance Cretaceous interval and a younger Pleistocene interval. Lancian is now applied
to the latest Cretaceous North American land mammal age and the Pleistocene is
divided into two land mammal ages, Irvingtonian and Rancholabrean, following Savage
(1951). All but one (Dragonian) of the 17 land mammal ages proposed in 1941, along
with the addition of Irvingtonian and Rancholabrean, are presently recognized by
North American vertebrate paleontologists. The currently recognized North American
land mammal ages are illustrated in figure 1. Dragonian was reduced to the status of
an earlier portion of the middle Paleocene Torrejonian land mammal age by Tomida
(1981). This was followed by Archibald et al. (1987) in the volume "Cenozoic Mammals
of North America" edited by Woodburne. Another North American land mammal age,
Clarkforkian, was later suggested the result of inaccurate or mixed stratigraphic data
for deposits that contain older Tiffanian and younger Wasatchian fossils (R.C. Wood,
1967). However, later intensive recollecting and biostratigraphic study in the type
area for Clarkforkian (and adjacent strata) in the Clark Fork Basin of Wyoming by
Gingerich and students demonstrated the reality of the Clarkforkian (Gingerich and
Rose, 1977).
Characterization of North American Land Mammal Ages
The Wood Committee Report characterized each North American land mammal
age by (1) a type fauna, commonly the accumulated fossil remains from some
restricted lithostratigraphic unit (e.g., Blancan is based on the local fauna at Mt.
Blanco and adjoining draws near the "old rock house," north of Crawfish Draw, Crosby
County, Texas), (Z) principal correlative faunas, (3) index fossils (considered restricted
to that land mammal age), (4) genera appearing in that land mammal age, (5) genera
last recorded from that land mammal age, and (6) genera characteristic of that land
mammal age. Thus, each land mammal age was defined on paleontologic data from an
identified lithostratigraphic unit and a geographically restricted area. However,
lithostratigraphic and geographic characterizations were not included.
North American land mammal ages are biochronological units, representing

spans of time during which the characterizing fauna lived. The principle data were
the occurrences of fossil mammal remains, representing taxa that lived near the site
of deposition at a specific time; conceptually, this represents the fossil equivalent of a
neontological "fauna." Some of these faunas have a firm biostratigraphic foundation,
such as the middle Eocene Bridger fauna, illustrated by Tedford (1970) and reproduced
here as figure 5 to emphasize the biostratigraphic basis for Osborn and Matthew's life
zone concept. Note the sequential appearance of brontotheres and Uintatherium, that
permit good chronologie resolution of this fauna. The term "fauna" has often been
606
Pol~osyops lorrtinotis
i.\:
- - - f* - ----l--f--+-1- +-+-+-+- 1--+--+- ~ Polaaosyops pol11dosus
f - - + - -- +-1.. ..-- - ~- r-- S' ~1--
-1-f--f·--t--L-,im_n_o_
hy'o--
'Ps-m
_o_n-
oc-o-
,.,.
---l
- - - -· ~ l: ~ 1--+-t- -f-....,-,,---:'----- - -1
~ _ ~ -- \ -- ~-1--+-t--t--L-im
_no
_nr_•_
'P•_m-,ot_
th_e"_;_---i
~ \ ----loL.- LimnO,)'I>PS priSC(IS
1----+-- ~ - -i~~
~+--+--+--~-4-4--~+-~~~--+-~-+--,_~~----~-----1
1--- -1-- _ i_.
_ -:-_!::. : --1---l--+-_..,_._,\·\1-·----+~::_:,:..:.. .:..:._:_ ~ ~-----+-·--+·---+----t-*---1--t-'..:?_JM_•_s_ot,_ih_inu_s_,"-_"'_u•,---;
Poloeos.tops major
""' ~-.. - -"'*- Polotosyops grong~ri

~-- - --~ -- --~ - -- - - -
~ )) ~ ~
---- Monfl!oceros manteocllt'tJS
- ~-- - 1-- - - ~ - ..., .. - . - ... :: - - - ---- -- - - - ---1
!---- -
r-- - -1--+-- ~ -t--• - -

~ r*+·*- ···f-*- -- -- ---
t::_ - ..- 1--- --- - -· ~
_!!JTetmothe1ium culllitltns
N"g - --
-- .. --! - 1---- \-~--=-~::: ~= ~ - _-- -- M•solilhinus pt/•rsoni
!----· - - - -- --- f- l --. : \

N ---~-0':--~~q~::j_-l---1--1-- - f?J~~<*>P<'!'_~::':!.!_
Pola~osyops Mldyi
r-- ~: 1: -- ---= - -- - - - - -- - - - -- 4
o ~_ ...-.. • - - - -
f- -- --- -~--~ a - -~
~ ~
1\_
Mesotirhlnus m'gorhlnus
- -- -- 6- -----· c......... _ Poto.o;;;;;;-;;;.,-- - - -
- f ___
.. a.·-
. . __ f4<- ["""'"""-
-·--· _r-
- --
- - - --- . _
1
~
"'*- . ..
. - f- - -
Poto•osyops 1o/JIJstus
-_!!_
-in=-to;_he
- _, iu=m
=-·-P-._-_-_-_-
__-_-
+----1f--f--+-~-+-+-+--lf-+-+ ..-..-_j-[----+. __ ___ __ -~.:..
oh....:'P::'.:.. •P_:_-- - - --i
P"•:...:.:
Fig. 5. Biostratigraphic ordering of large mammal appearances in the Bridger

Formation of Wyoming. (Figure Z in Tedford, 1970.) The actual
record of each species is indicated by the "X" and heavy vertical line;
gaps of nonoccurrence or unverified occurrence is indicated by dashed
vertical line.
used by vertebrate paleontologists for the remains from two or more sites (or local
faunas) that were more or less contemporaneous and shared most of the same taxa. In
paleontology the term "fauna" has acquired a temporal factor, a fourth dimension,
especially in the application to biostratigraphic data where segments of a faunal
sequence are also called "faunas" (see Tedford, 1970, for example). Wilson (1975) has
pointed out that in such cases faunas appear to represent biostratigraphic units in
another guise, but as he admits the present American Code of Stratigraphic
Nomenclature (1962) defines biostratigraphic units as bodies of rock characterized by
their fossil content, not as biochrons characterized by an assemblage of animals living
together in space and time (i.e., the paleontological connotation of the term
"fauna"), This emphasis on mammal occurrences in time rather than in rocks
distinguishes North American vertebrate chronology from other chronologie systems.
With subsequent collecting and taxonomic revision during the last 40 years the
fauna identified with each land mammal age has been revised but the criteria for each
of the land mammal ages has remained the same. It is a credit to the wisdom and
expertise of the paleontologists who compiled the data for the Wood Committee
Report that so much of it is still reliable.
607
A B c
TEXAS GULF COASTAL WESTERN OLD WORLD POTWAR PLATEAU,
PLAIN PAKISTAN
1- 8
z
w
en 9
w ,,
a:
/////////////~/-' //
fl. 10 ./9?~~~~i~9~r}~~ .-r1_~9r}~~s)~/

w
a: //////////////////////
0 11
LL •Hi~IH~riQn .. (CgrmQhiggariQn-
w ~like) "orimigenium"/~
m 12 //////////////// /
en
a:
< 13
w
> 14
z
Q
..l
..l
15 ~j~((;(((.(!!{((;{~~
//////////////// '/
:::E 16
Fig. 6. Sequential appearance of the horse "Hipparion" in North America,

Europe, and southern Asia. (Figure 3 in Flynn et al., 1984.)
Principles and Practices in North America
Another contribution to North American biochronology written by R.H. Tedford

(1970) was titled "Principles and Practices of Mammalian Geochronology in North
America." Tedford summarized practices and terminology applied to vertebrate
chronology from the time of its development until the time of his writing. Tedford
reviewed concepts and applications subsequent to the Wood Committee Report; he
clarified several terms and concepts, especially those in the Wood Committee
Report. Tedford (1970, p. 688-689) pointed out that North American land mammal
"ages" are neither equivalent to geochronologic ages (e.g., divisions of epochs) nor to
their equivalent chronostratigraphic stages. There was an implicit similarity of land
mammal ages to chronostratigraphic stages written in the definitions in the Wood
Committee Report (i.e., both are based on lithologic and chronologie or faunal data)
but the intention of the committee was to propose a series of sequential biochrons.
Savage (196Z) noted that land mammal ages lack the biostratigraphic-lithologic
foundation upon which chronostratigraphic stages are based. Both Savage (196Z) and
Tedford (1970) argued that vertebrate paleontologists should document the strati-
graphic record of mammals more accurately, to attach a biostratigraphic foundation
to land mammal ages and bring mammal chronology closer to the biostratigraphic
framework of marine stratigraphy. This recommendation has generally been
followed. Two of the original North American land mammal ages (Clarkforkian and
Wasatchian) were proposed as stages during the last 10 years (Savage, 1977; Rose,
1981). However, this has not resulted in greater resolution nor accuracy for those land
mammal ages. Biostratigraphic data acquired since 1941 has vastly improved the
degree of resolution exhibited by North American land mammal ages but their
calibration was acquired primarily through the application of radiometric and
paleomagnetic data to biostratigraphic data. Such data allowed Flynn et al. (1984) to
test for "heterochrony" (i.e., homotaxis) and to examine the temporal resolution of
land mammal ages (see figure 6). Biostratigraphic data always enhance land mammal
ages, especially when the biostratigraphic data become the framework for the addition
of other chronologie data.
The Woodburne Volume
The recent volume "Cenozoic Mammals of North America" edited by Woodburne

(1987), referred to below as the "Woodburne Volume," is the only comprehensive pub-
608
lished review of North American land mammal ages; each chapter dealing with a set
of land mammal ages (chapters 3-7 in Woodburne, 1987) builds on the data presented in
the Wood Committee Report, noting additions and revisions, to broaden and clarify the
characterization for each land mammal age and to ponder, for the first time, accurate
definitions for each age. It should be noted that the Woodburne Volume was a team
effort, similar to the Wood Committee Report, with 41 contributors, including 38
vertebrate paleontologists.
Boundaries of Land Mammal Ages
Probably the most significant problem in application of any mammal chronology

is the definition and recognition of the boundary between two units. Two faunal
assemblages in separate chronologie intervals that are close to the same boundary are
more easily united than separated. Thus, definition of boundaries between chronologie
intervals must be well defined and rigorously characterized. This problem was not
directly addressed by the Wood Committee Report and was only alluded to in
stratigraphic codes prior to about 1980. In recent stratigraphic codes the lower
boundary of chronologie units is always addressed, and is usually based on the
appearance of a new taxon or taxa. Overlying (and younger) chronologie boundaries
are usually not addressed, but are designated when the next (younger) chronologie unit
is defined. Recognition of upper boundaries of biostratigraphic units is not required
for their characterization. This practice has been widely applied by vertebrate pale-
ontologists in North America, often selecting the appearance of an immigrant to
define the boundary, as suggested by Repenning (1967) and elaborated by Woodburne
(1977).
However, selection of the earliest among several nearly contemporaneous

records of the fossil immigrant will always remain a problem. Most of the faunal
divisions and subdivisions identified in the Woodburne Volume are defined by the
appearances of immigrant taxa, and in many instances the specific site considered to
represent the earliest among several records of that immigrant is given. An attempt
to resolve the problem of "real" first appearances was provided by Lindsay et al.
(1987) in the Woodburne Volume. They distinguished "lowest stratigraphic datum" or
LSD from "first appearance datum" or FAD; the LSD represents the lowest known
record of that taxon in a stratigraphic sequence; the FAD represents an interpretation
of the earliest record of that taxon among its numerous fossil records in any biogeo-
graphic region.
Summation
In summary, North American land mammal ages are now well established and
confidently applied. The taxa that characterize and identify these land mammal ages
have changed as the fossil record became better known, but the principles upon which
the land mammal ages were established are still valid. Indeed, North American land
mammal ages now come very close to "representing all of Cenozoic time with little or
no overlap of included time." Within the context of scientific progress discussed by
Kuhn (1970), mammal chronology in North America has advanced beyond the observa-
tional level of fact-gathering. The framework initiated by Osborn and Matthew and
formalized by the Wood Committee Report has been tested during the last 40 years.
The Woodburne Volume represents a refined, broadened and articulated data base.
Further refinements will be needed and useful, but the main thrust and direction of
advances in the history of mammals in North America will more likely come through
applications of this data base.
EUROPEAN LAND MAMMAL FAUNAS
Traditionally, European mammal faunas have been placed in a framework of

European marine stages. This grew from (1) the initial development of the geologic
time scale within the framework of European marine stages, and (Z) the ease of
placing terrestrial deposits relative to marine deposits in the Paris Basin, a key strati-
609
EPOCH Ma MPTS MAMMAL AGE SUBAGE RECOGNITION
OR ZOt£
~
w
u T
0 1211
(!) CHADRONIAN
_J
0 T
40
OUCHESNEAN
Latt
45 UINTAN
w Earl'
z
w
u -- -'!!.'-I Epilfil/llfl#. ~ r,,.,.. F,.,,.,_,.,,., AmJ'IfiiiiiD\ ~ ,.,.,,,.,
0
w
1521
T T.-...utttltfl ="=•t'*•S. plus TWtrttiiiWillltf, ~ ._..,,,.._ ~ ,.e1111,
BRIDGERIAN .._.,_Ilion
-?-? -?- No/INNCII# Nll/l41it/l', Hottltlctlt/tJI ......... Oro/lil/lll4 ly/tlfllltd, , . ,....
r,.,....,
,_w,.
rl/loi/M, ....,....,.. ,.,,.,..,..,_ Ntllltflnt.n,.,.,.... S/Nitltlllt:lft f'flt:l"li.. OnJ/HtiiiW
'"J •w
- - - t»i
T _ i:
-T BN
.. _,_,_,_,
_ ~·-
_- - _ r=,._JifJIM.
llflllliiiW.
,.,.,,.,
~
r,.,._,
,,;,,..,. ~,.,.,. .,.,.,.
~ ,.,...,. ,.,_,., _,.. ._,_ -- .,_.
,.., .,.......,.., -.
I'·
~
LGStcallinlan
IV-II
I
.. N
c _ .. __ J~,.,..., ~. A"ti«<ttfJJt. OnJ/tillilfll, lhfti/Hiflo/lyll, ~ ScNnwn.
~;t,...., -..,.
IL Lfllteclll
WASATCHIAN
STAGE/AGE
=·
T :i ..-...;.;; liOhCirizon A, Ho#llogtl1t111 ~,.
------ -:Ji:-r- Cti"!'- f::.=ta£,.=·~.:-:C:: =~~ Sc,:;;7.'::bo,';:it:,.,

CL~~~9:iAN =:~~--- ,_._,-:';" .,,.._,,..
- - - - - - ~!~ J-: -;~~~- - =::.::..,.:j~=:C~i ,,.~ ,.,,. 1311
Ti5
60 T
Ti4
TIFFANIAN TiS
w
z Til
w - -?- -?- - -?- - -!i~', :Til;~·
u T T.S
0
w TORREJONIAN Toz
_J r.,.,.~t~UtH~tHt
~:~T;t
~ 65 --?--?--?-+~=~~a£"!.;~-
"-'•PI~INI ctNitrifttll
~ .. _ ;:{ _ _ ro.,~kiiJ,, ,...,,,
,~1------~·~''--!JI--111-111-.....,.f-:.11 ~==·
Fig. 7. North American land mammal ages and their subdivision. (Figure 10.1
in Woodburne, 1987.)
610
EPOCH Ma hf'TS MAMMAL Af£ RECOGNITION
E~~~~yt~rit,_,'-'. £.,.1/i..,.._ llfllltnfH. Ltllnl¥111. ..,_.,..,..._ NH/NifiiU'H, PlitltttlnfiN.

Sim«,-, TltilttJIJotl/sl• 1111
Note: Meplonyz sboald be basal Heaapbilliaa
-----'!!!--
LJ.J
z BARSTOVIAN
T
LJ.J
u T
0
:::!:
_ _ _ _ 1!!1 _ _
llltN:k~. c_,., Mitllllfnl•ltl. l#iollil:fis, Pliomefi., "-1t1Wistlldott, ~lw11..

. T•~Hunn 117'1
HEMINGFOROIAN
ARIKAREEAN
LJ.J
z
LJ.J
u
0
Q
...J WHITNEYAN T
0 _J.H]..--
'"'twt:"-'""· $ct11timus hlp/Nitus, Pnii«Vtls c•'-'· ,._,.,.,,.,, possibl,, t.IJ., Eumy1
I ORELLAN_
T /JI'tleii,Odfll, HytiMIOI/OIIIH.IIirOSiri.. Ni,,llf/1 IJNICIIytJPI, 0Jt.I0Cytll' CIISPUMIIIS 1261
Hyt/MtHifJn lloutdus, H. "IICH/111, lscllpomys, Eumy., l'tll'tldjid#lflllttJ lrilfllt/lfiS, Adjidoumtl

T MtiHI/11$, H. .~liCifHI gr•ptlll, ~. EulyiHII'fyS 1/klmpiOIIi, ~. thlo/11/11Jf4,
~IIIHklff/6 llfiJI#IMt, Hy~W'INifJfiiiiS 1271
611
Zonu A.gu g~ologiquu
PUistocble
Perplgnan P1ioc6ne s. str. N6og~ne sup6rieur

(Faunes 1\ Hipparion)
Teruel • PonUen •
Saba dell
La Grive N6og~ne moyen

Sansan (Mioc6ne p. p.)
La Romleu
Laugnac N6og~ne lnf6rieur

Paulhiac (Mioc6ne p. p.)
Coderet Ollgoc~ne sup6rieur

Co urnon
Antolngt
La Sauvetat 01igoc6ne moyen
Ronzon
Montmartre 01igoc6ne inf6rieur
Euzet Eoc~ne sup6rieur

Robiac
Castres
Issei Eoc~ne moyen
Argenton
Culs
MuUgny Eoc~ne lnf6rieur
Meudon
Cernay Pa16oc6ne
Fig. 8. Sequence of mammal zones for Tertiary mammal faunas

of Europe. (= Table 5 of Thaler, 1966.)
graphic-paleontologic reference sequence for (i) marine stages, (ii) the Cenozoic time
scale and (iii) vertebrate paleontology. The development of European Cenozoic
marine stages and their relationship to European mammal faunas was skillfully
reviewed by Savage and Russell (1983) in their global compilation of mammal faunas.
In fact, Savage and Russell (1983) use marine stages for the European Eocene mammal
faunas (except for Robiacian), whereas other contemporaneous references to European
Eocene mammal faunas (e.g., Fahlbusch, 1976) have generally adopted European
mammal ages or mammal stages for the mammal subdivisions.
Mammal Zonation by Thaler
Thaler (1966) recognized the futility and inherent complication of correlating

European continental deposits and faunas with the chronologie framework of marine
stages. In 196Z at a colloquium on the Paleogene held at Bordeaux, Thaler proposed a
halt to the correlation of European mammal faunas with marine stages, replacing
these with a sequence of European Tertiary mammal zones for the mammal faunas.
These mammal zones are illustrated in figure 8 (= table 5 in Thaler, 1966). Thaler had
recovered a number of new fossil small mammal assemblages utilizing acid
preparation and wet screening of sediments. In 1966 he described these faunal assem-
blages, collected in the vicinity of Montpellier in southern France, and placed these
faunas in a European mammal biozonation, based primarily on rodents (Thaler, 1966).
He proposed (Thaler, 1966, p. 187-189) Z3 mammal zones for the Tertiary of Europe.
His mammal zones were based on mammal assemblages from well known European
sites, and a type locality was designated for each zone.
612
Thaler distinguished these mammal zones by "stage of evolution" among
selected, well established lineages of mammals rather than by superposition. Most of
the reference faunas designated by Thaler had previously been correlated with marine
stages (e.g. in the Paris Basin), so the sequential ordering of these zones had already
been placed in a chronologie framework based on marine stages (e.g., Stehlin, 1909).
All of Thaler's mammal zones except one (Euzet) were defined on the basis of fossil
rodents, and all of the zones except two (Teruel and Sabadell) were located in
France. The fossil materials that Thaler studied and described (in 1966) were domi-
nantly Eocene and Oligocene; therefore, his characterization of mammal zones during
those epochs was more complete than for other epochs. Another important attribute
of Thaler's mammal zones is that his chronologie divisions were implictly shorter in
duration than the marine stages and thereby capable of greater chronologie resolution.
Thaler (1966) discussed the history of collecting and study of the reference
faunas (e.g., Cernay for the single Paleocene mammal zone; Meudon, Mutigny and Cuis
for early Eocene mammal zones, etc.) and rationale for the chronologie order that he
recognized. Stage of evolution was based on species differences in several well repre-
sented genera or lineages (e.g., paramyid, pseudosciurid and glirid rodent species for
the early Eocene zones; Lophiodon and other perissodactyla for the middle Eocene
zones; theridomyid rodent species for the late Eocene zones) to justify the recognition
and sequential ordering of zones. Where the faunal content was more complete, guide
fossils for specific zones or subzones were named (e.g., Isoptychus aquatilis for the
Ronzon zone), Thus, a sequential ordering of mammal faunas was established on the
basis of paleontological attributes (e.g., stage of evolution). Correlation with
European marine stages was not discussed although that information was available in
the literature for many of the fossil sites. This was a true biochronology, with chrono-
logie ordering based primarily on stage of evolution in selected fossils. On the other
hand, chronologie coverage in the mammal zones of Thaler (1966) was uneven, and
geographic distribution of his faunal assemblages covered only a small part of Europe.
A number of subsequent studies on European fossil mammals, primarily those

published by French paleontologists (e.g., Sudre, 1969, 1972; Hartenberger, 1969),
utilized and expanded the biozonation proposed by Thaler. Thaler (1972) published a
more extensive analysis of chronologie ordering, repeating many of the arguments
presented in 1966. Tables presented by Thaler (1972) are reproduced here as figure
9. Note that Thaler used the term "faunizone" (an interval between two faunal breaks
or coupures) which closely approximated the concept of "life zones" by Osborn and
Matthew (1909). He used the term "biozone" for an expanded concept of his mammal
zones in which each biozone is bounded by specific guide fossils (niveaux-reperes,
following Hartenberger, 1969). In other words, Thaler's biozone may be divided into
two biozones or sub-biozones by identifying a new niveau repere within an existing
biozone. Thaler (197Z) provided a new series of Z3 "biozones" and 3Z "sub-biozones,"
listing the type faunas bounding these zones. The type faunas were placed into a
chronologie framework based on marine stages. Eleven of his new "biozones" were the
reference faunas for the previous mammal zones; however, 8 of the Z3 "biozones"
were from countries other than France, thereby expanding the geographic range of his
biozonation. Thaler (197Z) was concerned that the biozonation he proposed should not
be extended over large geographic areas such as all of Europe. He feared that subtle
differences in stage of evolution of lineages that permit chronologie ordering might be
masked by an ecological overprint, and temporal gaps between suspected local immi-
gration or extinction events might produce an unrecognized reversal of chronologie
ordering.
Influence of Marine Biostratigraphy
Advances in the resolution of marine planktonic biostratigraphy during the

1960s, along with calibration of the Cenozoic time scale by increasing radiometric and
paleomagnetic data, spurred the chronologie resolution of marine deposits. Boundaries
of Cenozoic planktonic foraminiferal zones were being identified and calibrated as
"datum planes" that permitted a high-resolution global framework never previously
achieved in earth history. Note the similarity of "datum planes" and niveatz!t-reperes.
613
0)
....
~
Nn.a-t71* Nombnde CorNiatiolll del m-ux-t:vpe~

ZcmeiEq...,.
GraDde coupure pliiltocine -~
28. SIMIIS (LaDpedoc) 4 ViiJaflulc:bieD. Moldaftell
Zcme i Hlpptlrlon
Coupure ftll&ienne
22. TEaUEL (.......)
- 2 Turalim. PlailulciiD ?
---
21. SABADELL ( .......) 1 ValliMn. Tonollilm ?
Zone i AnelaltMrium 20. LA GRJVE (R.b&ae) 1 Tonollilm?
GraDde coupure JDioc8ne 19. SANSAN (.Aquitaine) 1 HeJftUeD ?
Zone aAmph,.., 18. LA R.OIIIEU (Aqda!De) 2 Bllldiplila ?
Coupure cbattieJme l'l. LAUGNAC (Aqda!De) 2 Aqul&aaial ? BllldiplieD ?
Zone i AntllllleotMrium 16. PAUUIIAC (.Aqui&lline) 1 ChatUm ? Aqultallilm ?
GraDde coupure ou,oeiae 1&. lbatENBAat (8m.e) 1 a.w.a?
14. BONJNGEN (SW..) 3 Cbattiell?
Zone i Anoplotlaerilma
13. ANToiNGT (Auverpe) 1 Cbattiell ? StampieD ?
Coupure anoplotherieane
12. MONTAI.BAN (Ea...-) 1 StampieD?
Zone i PaiRotMrium
11. HooGaursEL (~ue) 1 Tollpiell. Stampiell.
Coupure paliotberieDne 10. LA DEuUGE (~) 1 LaUorlleD ? TOIIp"ien ?
Zone i Hyracot,.,.,m 9. Buur (I.Dpecloc) 1 Butollilm ? Lattorflla ?
GraDde coupure byncotMrieune 8. Boaw: (Wipedoc) 1 ButoaieD. MuinM!en ?
Zone i Arctocyon '1. LlssJEu (JlbOae) 1 ButoaieD. Auveniea
6. EGEiwNGEN (Suiue) 1 Butollilm ? Lut6Uen ?
&. BouXWILLEil (Aiace) 1 LutetiiiL
4. GIIAUVES (B. puiliell) 1 Cllilin ? Lut.etim ?
3. AVENAY (B. pui&iea) 2 CWiieJL
2. DoRIIAAL (Belpque) 1 Spunaciea.
1. CUNAY (B. puiliell) 1 Tbauetiea.
----
Fig 9. European mammal faunizones (left side) and biozones (right side) proposed for Tertiary mammal
faunas of western Europe by Thaler (197Z).
European Terrestrial •Biostratigraphy"
Vertebrate paleontologists in Europe were concerned that resolution in

mammalian chronology should keep pace with advances in planktonic biostratigraphy.
They were also concerned that European mammal chronology was tied rather inse-
curely to marine biostratigraphy that was being revised at a rapid pace. And, fear
that European mammal chronology would become chaotic if each nation were to
develop a "national" chronologie framework, like the almost exclusively French bio-
zonation of Thaler, stimulated vertebrate paleontologists in Europe to establish a
European terrestrial biostratigraphic framework for mammal history.
Productive Neogene fossil deposits were being developed in Spain during the
1960s and Crusafont (1951) had proposed a Vallesian "Stage" for faunas in the Valles-
Penedes area of Spain, to establish a Mediterranean or Tethys faunal reference
comparable to the Pontian" or Paratethys faunal reference that had been applied
throughout Europe for that late Miocene-early Pliocene interval. Later, Crusafont
(1965) proposed a Turolian "Stage" for faunas in the Calatayud-Teruel area of Spain,
recognized as younger than Vallesian on the basis of faunal change. However, both of
Crusafont's "stages" lacked the descriptive biostratigraphic foundation needed for
establishment of chronostratigraphic units. At the 1966 meeting of the Regional
Committee on Mediterranean Neogene Stratigraphy (RCMNS) held in Bologna these
problems were discussed and de Bruijn was asked to develop a biostratigraphic founda-
tion for the Spanish Vallesian and Turolian "Stages."
A Ruscinian "Stage" was proposed at the 1971 RCMNS meeting in Lyon, to

represent faunas older than Villafranchian "Stage" and younger than Turolian "Stage."
Reference faunas for Ruscinian were primarily from Sete and Serrat d'en Vacquer in
the Roussillon Basin of southern France. In part, Ruscinian was a replacement for
Csarnotan "Stage" proposed by Kretzoi in 196Z for equivalent faunas in the Pannonian
Basin of southeastern Europe. Ruscinian is also interpreted as the marine transgres-
sion into the Mediterranean area coincident with the beginning of the Pliocene
Epoch. Villafranchian "Stage" was proposed by Pareto (1865) for fossil mammals from
Villafranca d'Asti in the Po Valley of Italy. Villafranchian has been widely recognized
as a terrestrial equivalent to the marine Calabrian Stage in southern Italy, which
marks the beginning of the Pleistocene Epoch. Thus, a number of faunal units with
varying degrees of characterization had been proposed, usually as chronostratigraphic
stages, for late Cenozoic mammal units of Europe.
Berggren and Van Couvering (1974) summarized the status of European mammal
chronology, noting the disparity of a uniform set of stratigraphic rules and principles.
It should be pointed out that none of the "stages" noted above had been defined nor
described with the biostratigraphic foundation of a marine stage; technically, they
were neither biostratigraphic nor chronostratigraphic units. They were biochronologic
units, based primarily on changes in mammal history. Berggren and Van Couvering
(1974, p. 91-9Z) identified nine biologic and geologic events that mark significant
"breaks" in the late Neogene history of European mammals. Many of these "breaks"
are still valid, and have been incorporated into the emerging European mammal
chronology.
In 1976 A. van de Weerd, a student of H. de Bruijn, described rodent faunas from

the Teruel-Alfambra region of Spain, placing them in a biostratigraphic framework
with recognition of six biozones that span upper Vallesian, Turolian, and Ruscinian
"Stages" (see figure 10). Van de Weerd provided the biostratigraphic definition and
characterization for these biozones, but, more important, he demonstrated super-
position of previously-named mammal units; e.g., Vallesian, Turolian, and Ruscinian.
In reflection, the only valid chronostratigraphic stage developed by van de Weerd is
Turolian Stage. Van de Weerd characterized the position of Turolian relative to
Vallesian and Ruscinian, based on biostratigraphic data, but his study did not include
the earliest Vallesian, nor did he have fossils that permit recognition of the Turolian/
Ruscinian transition.
615
Selected localities Correlation with loc.lities outside Zonation Zonation
in the Teruel Biozones Thaler Mein Stag. . Series
Alfambra region the Teruel. Alfal!lbn region IIIIII 111751
Csarnou 2 16 c:
-----
Mimom.,, Moreda
Escorihuela -~
at•hlini
0
....:
BlllfUC 2
c:
•c:
·a."'
Silte Weze u
.--
•c:
Layna
Goraffe 2 •
...
...•
15
c: •u
0
."
c: ;;:
Orrios
Casrillomv•
CTUIIfOnti
Goraffe 1
laneue Serrat
d'en Vacquer
0
N
•c: ----- u
gracilia
a:
14
------ ----- -----

...•
CltiVICI
Valdecebro J
,.
•c:
Crev•llente 6 0
Stephanomrs !:! 13
.
La Fanran1 <(
ramblansil Chomater• 0
Masada del Valle 7 N
Polgardi
------ c:
Masada del Valle 5
Los Mansuetos
Paragodamus Crevillente 5 .. ·-0
•c:
...;
fiUt#'li 12
~
Concud 3
Mesada del Valle 2
barbaraa Creve I lente •
...• •u
Torujada A
Eichtr.ogel ...• 0
:1
Parapot1amu1
lugdunanlil
Mallon
Crevillente 1.2.3
Oorn. Ourckheim
•c:
0
11
Alfambra N
KohfidiiCh
------
Peralejoa C.O Progonomy1 Sob loy
...• ... 10. c:
•
••
.-
hi•p•nicu•
Masri del Barbo 21 Moncredon
•c: ...•
0
N • -----
Ill
>
C•n Llob•cerea 9
Fig. 10. Biozonation and correlation of selected mammal faunas in the Teruel-
Alfambra area of Spain by A. van de Weerd. (Figure Z9 in van de
Weerd, 1976.)
European Mammal Neogene, or MN, Zones
The need for a comprehensive European framework for mammal history was
apparent. P. Mein compiled fossil mammal data from Neogene sites in 14 countries
throughout Europe (i.e., Spain, Portugal, France, Germany, Switzerland, Italy, Austria,
Hungary, Czechoslovakia, Poland, Romania, U.S.S.R., Greece and Turkey) and north
Africa, ordering those mammal faunas sequentially as a series of biochrons, similar in
principle to the "biozones" of Thaler. Mein presented his chart in 1974 at an inter-
national colloquium on continental biostratigraphy at Montpellier and Madrid (see
figure 11). The initial chart listed 190 mammal sites or faunas, including 39 from
France (8 of which were among Thaler's 197Z "biozones" or their equivalent). Mein
organized these mammal sites into 16 zones termed MN (= Mammal Neogene) zones,
plus an earlier Paleogene zone (MN 0), and a younger Quaternary zone (MN Q1).
Mein's MN zones listed both large and small mammals, including (1) characterizing
species of well established evolutionary lineages (e.g., used for interpretation of stage
of evolution), (Z) important genera appearing in each zone, and (3) distinctive associa-
tions of genera that help to identify each zone. Figure 11 shows only the western
European faunas listed by Mein (1979); he also listed faunas from eastern Europe,
north Africa, and Turkey.
These zones were expanded and formally presented the next year at the RCMNS
meeting in Bratislava (Mein, 1975). The updated chart listed ZZ1 mammal sites and
included Yugoslavia in addition to the countries utilized in the previous chart. Many
more characterizing species, generic first appearances, and generic associations were
616
GERMANY
MN SPAIN FRANCE SWITZERLAND
zone PORTUGAL ITALY
Q1 La Puebla de Saint Vallier Schernfeld
Valverde
Coupet
Roccaneyra
Villaroya Villafranca
16 Seynes . BalanJC 2
Medas Etouaires, Trevoux
Vialette
Layna Sete Gundersheim 1

15
Escorihuela Perpignan Wolfersheim
Montpellier
14 Goraffe 2 Hautimagne
Hauterives
Alcoy
13 La Alberca Lissieu Casino
Arquillo Luberon
12 Los Mansuetos Ratavoux
11 Aspe Mullon , Lobrieu
10 Masia del Barbo Soblay

Villadecaballs Montredon
Can Llobateres Howenegg

9 Pedregueras St. Jean de
Nombrevilla Bournay Eppelsheim
Barbera La Grive 3 Anwil

8 San Quirze Giggenhausen
Can Mala 1 Oggenhof
Azambujera inf. St. Gaudens
7 La Grive M
Simorre
6 Manchones
Arroyo del Val Sans an Sandelshausen
Munebregs Pont Levoy

5 Lasplanes Langenmoosen
Sos
Valderamos 3 Suevres
4b Vieux Collonges
Baigneaux
4a Bunos Rubi La Romieu Ekertshofen

Artenay
Rubielos de Mora Chilleurs
3 Moli. Calopa Wintershof-West
Ateca 1 Estrepouy
Lisboa (Univ . ) Chitenay Bissingen
2b Bouzigues Haslach
Celina de Aragon Laugnac La Chaux
2a Marcoin Ulm
"St. Gerand" Budenheim
Saulcet Aarberg
Paulhiac Weissenburg 6
Weissenau
Boudry
Wischberg
Tomerdingen
0 Coderet Kuttingen
Hochheim
Fig. 11. Simplified chart of Mammal Neogene zones,

presented by P. Mein (1979), showing only the
western European fauhas.
617
added. The MN zones were revised again at the next meeting of RCMNS at Athens (in
1979), with some new divisions (e.g., MN 3a/3b and MN 16a/16b), revisions (e.g.,
MN 17 for MN Q1), additions (e.g., Terrats, Montopoli and Rudabanya) with corre-
sponding revisions and additions in the species lists. At the 1979 Athens RCMNS
meeting, Mein summarized tentative correlations of MN zones with the calibrated
Neogene planktonic foraminifera zones. Other conclusions Mein presented at that
meeting were (1) MN zones are not of equal duration, and (Z) evolutionary diversifica-
tion of mammals within Europe was probably minimal because of the size of Europe.
Mein inferred that many of the large mammals in the European fossil record were
probably immigrants from adjacent areas, such as Asia and Africa (Mein, 1979). A
further revision of MN zones is presented by Mein (Chapter 6) in this volume.
This biochronological framework (European MN zones) will continue to grow and

be revised as new discoveries are made; the data base will become more stable with
time. However, a problem with use of European MN zones, as with any relative
chronologie framework, is that it cannot be calibrated unless tied to some other
independent chronologie framework.
The M1Dlchen Symposium
Another significant step in the advancement of European mammal biochronology

was the symposium held at Munchen during April 1975 to consider principles and
formal designation of a European continental chronologie framework. The need for a
uniform and inclusive chronologie framework for European mammal faunas was widely
recognized. Thirty vertebrate paleontologists from six European countries attended
the Munchen Symposium. Fahlbusch (1976) reported that discussions were open and
intense; he listed three areas of concern and conflict: whether European mammal
faunas should be placed in (1) a biostratigraphic framework, with type sections and
formal characterization, as in marine deposits; or (Z) a biochronologic framework,
such as immigrations, evolutionary steps, and faunal change; or (3) in an arbitrary
framework, but with exactly defined boundaries, the lower boundary of each time unit
defined by an appropriate fossil locality. It was generally agreed that a sequential
ordering of mammal faunas had already been established, based on (i) previous
correlations with marine stages and (ii) evolutionary stage of evolution in well-
established mammal lineages. Identification of boundaries between land mammal ages
was discussed at length, but criteria for recognizing boundaries could not be agreed
on.
A compromise solution at the Munchen Symposium resulted in the selection of

reference localities from existing data, considering (a) homogeneity, (b) diversity,
(c) other paleontological content, (d) abundance, (e) evolutionary level, (f) minimal
ecological variation, (g) published knowledge, and (h) availability of the fauna. Forty-
five reference localities were selected, and grouped into 13 mammal ages (see figure
1Z). Reference localities for the Neogene had already been placed within MN zones,
so the Neogene European land mammal ages· had a sequential organization. Where
appropriate, faunal events would serve as boundaries between land mammal ages, and
for convenience references to previous marine stage correlations were given.
Characterizing assemblages for the land mammal ages would consist of the faunal lists
of designated reference localities. The resulting biochronologic framework, based on
mammal evolution in Europe, provided the possibility of refinement, revision and
precise boundary definition, as needed.
Aragonian Stage/Superstage
To illustrate growth and application within this chronologie framework, the

history of Aragonian Stage is useful. At the .1975 Munchen Symposium three "super-
stages" (Occitanium, Aragonium, and Catalonium) were designated as possible demon-
strations of European continental biostratigraphy. Aragonian Stage (or Superstage)
was identified as the interval between the appearance of the low crowned horse
Anchitherium and the high crowned horse Hipparion, comprising two land mammal
ages, Orleanian and Astaracian, approximately equal to early and middle Miocene.
618
I.,.JE-Iii------0 e f i n i t i o n-----~ ~~lnterpretati on~l
I! Reference Localities Mammal Ages Faunal Events Prtvioua Subdivision
H
"Superstates"
lrnp. Stagnl
NM17
AUophaiomys- Faunas
Villany 3
Blharium
--------
Pltistac:ent
Villanyium
NM 16 Rtbielice
JIM IS Perpignan Pliocene
Ruscinium
NM 14 Podlesice ,
NM 13 Arquilla /
NM 12 Los Mansuetos Turolium

/
/ ""
NM 11 Crevillente 3 Co tolonium /
NMIO Masia del Barbo 28
Vallesium / ""
""
NM 9 Can Llobateres
4-Hipptirion
NM 8 Anwil
NM 7 Steinheim Astargcium
NM 6 Sanson Miocene
Araggnium
NM 5 Las Planas 48
NM 4 La Romieu Orleanium
Burdigalium
NM 3 Wintershof- West
NM 2b Laugnac
NM 2a Montaigu !_g.!!!l!l!!! Aqultanium
NM 1 Paulhiac
I
Coderet I
La Milloque Chattium /
Arvernium
Boningen
Antoingt
I Oligocene
Occitanium _....{
Heimersheim
- -- II Stampklm
Montalban I
Suevium 1
. Villebramar Rupelium
Hoogbutsel ____ _lI___ _

Grando Caupure I
Frohnstetten I
Montmartre I
Lattorfium I
I
--- -- --
La Debruge
Perri ere
Headonium - - - - - - -;;:Jill------'
Fons 4
Grisolle
Robiac Bartonlum
Lo Liviniere
Lissieu
Egerkingen r Eocene
Rhenanium
Bouxwiller
Messel
Mas de Gimel
Cuisium
Grauves
Ave nay
Mutigny Neustrium Sparnacium
Dormaal
Cernay
Paleocene
Mons
Fig. lZ. Subdivision of European Tertiary mammal faunas, presented by V.

Fahlbusch (1976). Note the Neogene Mammal zones are placed to the
left of the principal reference faunas.
619
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Freudenthal, 1988.)
Aragonian Stage was formally established and characterized (Daams et al., 1977) for
sediments and fossils in the Calatayud-Teruel Basin of central Spain. Subsequent work
(Daams and Freudenthal, 1981) served to better characterize the biostratigraphy but
revealed that the lower boundary (the appearance of Anchitherium) could not be
identified with confidence in the type area. This work had produced a large sample of
small mammals on which nine biostratigraphic zones (A through I) could be recognized
within both Aragonian and lower Vallesian Stages. In addition, Daams and Freudenthal
(1981) found that (1) bipartite separation of Aragonian (into Orleanian and Astaracian
land mammal ages) was impractical and suggested that these two units be replaced by
a threefold division of Aragonian zones (lower, middle, and upper) in the type area;
and (Z) MN zones in the type Aragonian area may be diachronous. They further
pointed to several problems in application of small mammal faunas in Spain with MN
zones 3-5. Specifically, (1) a previously unrecognized interval of time may occur in
zone MN 3, recording the absence or near absence of cricetid rodents (e.g., the
cricetid vacuum); (Z) La Romieu, reference locality for zone MN 4a may be younger
than Vieux Collonges, reference locality for zone MN 4b, based on species of
Megacricetodon; and (3) rodent species from Pontlevoy-Thenay, reference fauna for
zone MN 5, closely resemble or may be identical with species from Vieux Collonges,
reference fauna for zone MN 4b. Daams and Freudenthal (1981) also noted the need
for complete and updated faunal lists for all the reference faunas of both MN zones
and European land mammal ages.
Ramblian Stage
Subsequently, Daams et al. (1987) have redefined the base of Aragonian Stage,
on the appearance of modern cricetid rodents, and named the Ramblian Stage for
strata and fauna underlying Aragonian deposits (see figure 13). This revision places
zone A of their earlier study (Daams and Freudenthal, 1981) in Ramblian Stage.
Justification for this revision was recognition that Anchitherium is too poorly repre-
sented in these deposits to serve as a biostratigraphic guide fossil. Daams et al. (1987)
identified the base of their Rambliaii Stage with the extinction of the Rhodanomys/
Ritteneria group of eomyid rodents; they did not indicate where this extinction occurs
stratigraphically. They pointed to problems in correlating Ramblian Stage with
Agenian Age and MN zones Z-3. They suggested Ramblian Stage is probably equiva-
lent to part of Agenian mammal age. Chapter 4 by Daams and Freudenthal in this
volume reviews the further development and history of Ramblian Stage.
CONCLUSIONS
In summary, Neogene mammal faunas of Europe have been placed in both a

biochronologic and a biostratigraphic frame of reference. Also, it appears that many
European vertebrate paleontologists feel that the biochronologic frame of reference
should eventually be replaced by the biostratigraphic framework when adequate docu-
mentation becomes available. As previously demonstrated in North America, this is
neither necessary nor practical. The biochronologic framework for European Neogene
mammal faunas should prove an adequate and .reliable chronologie system; however, it
should be subject to testing and modification, as needed. The biostratigraphic frame-
work for European Neogene mammal faunas can be incorporated into the biochrono-
logic framework; it has an essential role in both testing and calibration of the biochro-
nologic framework. Biostratigraphic applications will prove valuable in calibrating the
European land mammal ages, and in demonstrating weaknesses or discrepancies of
biochronologic correlations, as already pointed out by Daams and Freudenthal (1981).
In the context of scientific progress presented by Kuhn (1970), mammal chronology in
Europe remains in the observational level of fact-gathering. Vertebrate paleontolo-
gists in North America had tested those land mammal ages for more than 40 years in
order to develop the confidence and reliability that framework now enjoys. Only
recently (e.g., Archibald et al., in the Woodburne Volume) has it been possible to
demonstrate in North America the application of biostratigraphic data to show the
"equivalence" of mammal biochrons and chronostratigraphic units.
621
The history of mammals is known much better now than it was 80 years ago,
when Osborn and Matthew framed a North American mammal history in a continental
context. It has taken three generations of vertebrate paleontologists to achieve the
level of resolution and accuracy that North American land mammal ages now enjoy.
On the other hand, European mammal chronology has been separated from marine
stratigraphy for only about 15 years. Development of mammal biochronology in
Europe, independent from marine stratigraphy, spans only one generation of verte-
brate paleontologists. Progress is likely to come much faster in resolution and
accuracy of European mammal biochronology, partly because of improvedmethods and
fossil records and partly because of a better global model for mammal biochronology.
By the year ZOOO biochronologic systems in both North America and Europe should be
comparable in resolution and accuracy.
It should be emphasized that vertebrate paleontologists have chosen a method of

faunal reconstruction that lies outside the methods governed by stratigraphic codes;
their focus is on biological events rather than subdivision of the rock column. The
units in this historical reconstruction are primarily biological entities, faunas and local
faunas, rather than biostratigraphic units that are founded on rock columns. These
samples of evolving biota are ordered in time by reference to the stratigraphic record
wherever possible; however, direct observation of temporal relationships is rarely
available in fossiliferous continental deposits. This is especially true in fissure and
cave deposits that often contribute abundant well preserved fossil remains. In such
instances, stage of evolution may be the only method that yields an hypothesis of
temporal position.
Independent geochronologic techniques, such as isotopic dating and magnetostra-

tigraphy, can be used productively to test hypotheses of temporal position and for
correlation of faunas; they might also be applied to assess the age of fissure and cave
deposits. In conjunction with fossils, these techniques can be useful for the calibration
of vertebrate evolution; however, by themselves, these techniques can never establish
a system of biochrons. That task must rest with the faunal record.
Mammalian biochronological units, or mammal ages, are now being developed on

all continents. This reflects favorably on the widespread recognition of the utility of
the life zone method of Osborn and Matthew in which biochrons are identified and
characterized from the faunal sequence. By reference to favorable situations where
biostratigraphic relationships of these faunas are fully depicted, it should be possible
under these constraints to transform the purely temporal scale into a chronostrati-
graphic one. Biochronologic and biostratigraphic methods for marking geological time
are not antithetical but are complementary, and each may have special utility under
different conditions. The nature of the vertebrate fossil record in terrestrial deposits
generally favors application of the biochronologic method.
Principles and practices of biochronology were recently discussed by Woodburne

(1987); these discussions serve as a valuable guide to the use of this method. More to
the point, we recommend that biochronologic units should be fully characterized; that
is, with designation of (a) reference faunas, (b) restricted species, (c) association of
genera, and (d) genera appearing in each unit. Criteria for recognition of boundaries
between adjacent units should be defined, whether the units are biochronologic or bio-
stratigraphic. Finally, complete faunal lists of principle reference faunas is essential
for the successful application of the biochronologic method.
REFERENCES
Archibald, J.D., Clemens, W.A., Gingerich, P.D., Krause, D.W., Lindsay, E.H., and
Rose, K.D., 1987. First North American land mammal ages of the Cenozoic Era,
in Woodburne, M.O. (ed.), "Cenozoic Mammal Faunas of North America," p.
Z4-76.
Barry, J.C., Lindsay, E.H., and Jacobs, L.L., 198Z. A biostratigraphic zonation of the
geography, Palaeoclimatology, Palaeol!'cology, v. 37, p. 95-130.
622
Berggren, W.A. and Van Couvering, J.A., 1974. The late Neogene, biostratigraphy,
geochronology, and paleoclimatology of the last 15 million years in marine and
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624
THE PAST, THE PRESENT, AND THE FUTURE
Volker Fahlbusch
lnstitut fUr Palaontologie und historische Geologie

Richard-Wagner-Str. 10
8000 Miinchen Z, Federal Republic of Germany
Pierre Mein
Departement des Sciences de la Terre

Z7-43 Bd. du 11 novembre
696ZZ Villeurbanne Cedex, France
Biochronology and biostratigraphy on the basis of European Tertiary mammals

were the subjects of a symposium held in Munich in 1975. Stratigraphers had early
recognized that active Neogene tectonism caused Europe to be separated into many
isolated sedimentary basins, which in turn caused discontinuities in mammalian distri-
bution and evolution. These very discontinuities were seen as potentially confounding
attempts to correlate mammalian localities. As a group, the symposium participants
departed from the traditional practice of relating fossil mammal faunal units to
marine stages, by proposing independent continental chronologie units. In following
years these units were related to marine stages only when there was a physical strati-
graphic interdigitation of marine and continental rocks. This practice paved the way
to remedy long standing confusion about the age and correlations of Neogene mammal
faunas. The newly emergent correlation system has proved quite useful for inte-
grating the tremendous volume of new data from many European countries.
The Munich symposium dealt with a new concept presented by Mein (1975) for
subdividing the European Neogene on the basis of mammals. This concept was an
extension of a practice developed during the last century whereby the time correlation
of more important mammal localities in Europe was based upon three principal
criteria: (i) the stage-of-evolution of individual species, (ii) their species associations,
and (iii) biogeographic aspects. Localities then were biochronologically ranked by
Mein into MN units. Participants in the Munich Symposium vehemently debated the
theoretical background of this zonation, the possibility of having well defined bound-
aries separating the MN units, and the proposal to combine two or three contiguous
units into larger units corresponding in time to stages or ages. There were some terms
already in existence for these larger units: Vallesian (for Mein's 1975 MN 9-10) and
Turolian (MN 11-13). Also, some new units were proposed: Agenian (MN 1-Z),
Orleanian (MN 3-5), Astaracian (MN 6-8). Furthermore, the symposium participants
proposed the erection of two "superstages/superages," the Aragonian (Orleanian and
Astaracian combined) and Catalonian (Vallesian and Turolian combined) (see
Fahlbusch, 1976).
In spite of some fundamental objections to the subdivision proposed by Mein and

the Munich symposium, and despite serious doubts about its practicality, this system
has proven rather useful and was accepted by many colleagues within and outside of
Europe, including not only mammalian paleontologists. To a certain degree, it turned

out to be applicable even for long distance correlations. Therefore, a somewhat
refined edition of the stratigraphic chart is presented in this volume (Steininger et al.,
Chapter Z), containing the correlation with the Tethys and Paratethys stages, as well
as other subdivisions. In the MN zonation a further subdivision (a, b) has deliberately
been renounced in most of the cases because in far-reaching correlations an accuracy
may be asserted which does not really exist. The stratigraphic chart and the updated
MN zonation by Mein (Chapter 6) contain a number of new data, but principally this
zonation is unchanged and corroborates the usefulness of this system of subdivision.
Meanwhile, a comparable subdivision has been proposed for the European Paleogene
(Schmidt-Kittler, Chapter 3).
During the Munich symposium in 1975 it was pointed out that a very important
subject of further study should be the intensive invesUgation of complete sections of
mammal-bearing terrestrial sediments, as far as those are available. On that score,
the last years have yielded remarkable results. By far, the most important area in
that respect turned out to be Spain (see e.g. Daams and Freudenthal, Chapter 4).
There it was possible, too, to establish biostratigraphic stages according to the Inter-
national Stratigraphic Guide, and additional stages are expected in the near future,
not only for the Neogene. Similar sequences of mammal-bearing sediments have been
investigated in Switzerland and Bavaria. In addition, from some of these areas there
are stratigraphic results available from other groups of organisms. The possibilities
for application of independent chronologie methods in those areas do not seem to be
exhausted. This is true even for magnetostratigraphy, a method which has been
proven to be very useful during the last years. However, due to the paucity of
complete sections with mammals in Europe, little attention has previously been paid
to this method. The papers presented during the Reisensburg meeting on that subject
(Chapters Z9-31) clearly demonstrate the application and results to be expected.
The investigation of complete sections will be very important in the future

also. It will result in the establishment of local sequences of biostratigraphic units.
However, in order to reduce confusion in our communication by broadening the appli-
cation and/or creating new stratigraphic terminology, it may be necessary to maintain
a stable reference system which is applicable for all of Europe. The participants of
the Reisensburg workshop agreed that, for the time being, this purpose is well served
by the MN zonation.
There is no doubt about the utility of documenting detailed biostratigraphic
investigations within continuous sections of limited areas. Only these investigations
yield reliable data for the resolution of biogeographic problems. Without sufficient
knowledge of faunal sequences in limited bioprovinces, it seems highly premature to
consider discrete migration events as well as extinctions or disappearances of single
taxa to be isochronous, and in consequence to use them as time markers. This is best
revealed by the immigration events (Proboscidea, Hipparion, Equus) which have been
under discussion for many years, and have been dealt with during the Reisensburg
symposium (see Chapters 16-ZO). It also becomes apparent in light of new results
applying the record of micromammals from Greece and Turkey (e.g., the missing of a
"cricetid vacuum" in the Lower Miocene in comparison with central and western
Europe). The precise temporal correlation of the Greek and Turkish localities with
those from central and western Europe still remains problematical in detail. This, on
the one hand, may be dependent on major biogeographic differences which are docu-
mented by distinctly separate faunal associations as well as faunal sequences. On the
other hand, there may be a point reached (at least with rodents) beyond which a
comparison on the basis of stage-of-evolution is only conditionally possible as long as
there are only locally and/or temporally limited lineages available. Judging from the
last years' results, these gaps in our knowledge may soon be filled. Anyway, without
additional data from adjacent areas, many details in the history of European mammals
will remain unknown.
A trend seen for many years in mammalian paleontology in Europe - as in other
626
countries as well - became apparent during the Reisensburg meeting. That is the
dominant role of micromammals, and rodents especially in the study of mammal
chronology. The large gaps in knowledge which existed for a long time in the history
of these mammals have been filled remarkably. This results primarily from the possi-
bility of dealing with larger numbers of individuals from many localities. However,
there are still tremendous gaps in our knowledge, and much effort will have to be
made in the research on micromamals in Europe, and even more in adjacent areas.
In spite of all the remarkable results which have been contributed by the aid of
micromammals to biochronology, biogeography, and evolutionary problems, one has to
consider as well whether more attention should be paid in the future to larger
mammals. Their importance became apparent just by dealing with biogeographic
processes and migration events during the Reisensburg meeting. And they seem to be
especially important from the paleoecologic point of view. More detailed investi-
gations on different mammal groups, especially artiodactyla, seem necessary and full
of promise for the future.
Furthermore, it seems worth mentioning that during the NATO workshop

comparatively little attention was paid to phylogenetic systematics, although cladistic
methods surely demonstrated better possibilities to elaborate relationships between
related forms which may often be hidden under too many generic names. This again
may be especially true with rodents. True relationships of rodents are quite often
doubtful because of their high diversity and many parallelisms in tooth structure (in
many cases nothing else is known); their systematics is uncertain and may still contain
many paraphyletic groups. In addition to a continual need to increase our data base,
more attention should be paid to these problems in the future if the methods of phylo-
genetic systematics are applicable to the remains of micromammal dentitions.
One subject which was addressed only slightly during the Reisensburg symposium
is paleoecological analysis based on mammals. Although this type of research is
actually done at many places in Europe, the organizers of this meeting intentionally
suppressed presentations of this subject in many instances. This did not mean an
underestimation of its importance. On the contrary, they agreed that paleoecologic
analysis is too complex to be treated without including paleobotanic data and that of
other specialists. Paleoecological problems of terrestrial fossil associations, and
paleoclimatic interpretations based on them, might be addressed better as the main
topic of another special meeting, with specialists representing several other disci-
plines also contributing data and discussion. This became evident during the
Reisensburg discussions related to those problems (see Chapters 25-28) as well as with
such phenomena as differences in faunal associations of stratified localities and
fissure fillings in western Europe. Another aspect was elaborated on by Andrews (see
Chapter 28). He clearly demonstrated the misinterpretations that are possible in
fossil associations (especially with micromammals) by taphonomic processes, and
which misinterpretations are possible in respect to paleoecology. Nevertheless, paleo-
ecology must be a subject addressed by European mammalian paleontologists in the
future.
Only little attention was paid to global events, e.g., sea level changes and their
influences on mammalian history. The need for more research in this respect became
clear, and this aspect of mammalian history must also be addressed in the future.
In summary, it may be stated that during the Reisenburg symposium a useful

documentation was presented on the results in European Neogene mammal chronology
during the last years. Many of these contributions have been published in this
volume. Not less profitable for the participants was the opportunity to learn from the
extensive (and sometimes controversial) discussions and conversations, those major
points of agreement and/or disagreement, as well as those goals and trends to which
mammalian paleontology and paleontologists in the Neogene of Europe and adjacent
areas should be directed in the future.
627
REFERENCES
Fahlbusch, V., 1976. Report on the International Symposium on Mammal Stratigraphy

of the European Tertiary. Newsletter Stratigraphy, v. 5, p. 160-167.
628
CONTRIBUTORS
J. Agusti B. Engesser K. Kowalski

Instituto de Paleontologia Naturhistorisches Museum Polish Academy Sciences
Sabadell, Spain Basel, Switzerland Krakow, Poland
B. Alpagut V. Fahlbusch J. I. Lacomba
M.T .A. Enstitusu Universitat MWlchen Universidad de Valencia
Ankara, Turkey MWlchen, Germany Valencia, Spain
M.A. Alvarez-Sierra 0. Fejfar C. G. Langereis
Universidad ·complutense Ustredni Ustav Geologicky University of Utrecht
Madrid, Spain Prague, Czechoslovakia Utrecht, Netherlands
P. Andrews L. Flynn E. Lindsay
British Museum (NH) Harvard University University of Arizona
London, England Cambridge, MA, U.S.A. Tucson, AZ, U.S.A
M. Antunes M. Freudenthal N. Lopez Martinez
Universidade Nova Lisboa Rijksmuseum Geol. Miner. Universidad Complutense
Lisboa, Portugal Leiden, Netherlands Madrid, Spain
A. Azzaroli L. Ginsburg F. Masini
Universita di Firenze Mus. Nat. Hist. Naturelle Universita di Firenze
Firenze, Italy Paris, France Firenze, Italy
J. Barry C. Guerin P.Mein
Harvard University Universite Claude Bernard Universite Claude Bernard
Cambridge, MA, U.S.A. Lyon, France Lyon, France
R. L. Bernor M. Gurbuz E. Moissenet
Howard University M.T.A. Genel Mudurlugu Universite de Paris I
Washington, D.C., U.S.A. Ankara, Turkey Paris, France
M. Bonifay W. Heinrich S. Moya-Sola
Lab. Geologie du Quater. Humboldt University Instituto de Paleontologia
Marseille, France Berlin, Germany Sabadell, Spain
H. de Bruijn K. Heissig N. Opdyke
University of Utrecht Universitat MWlchen University of Florida
Utrecht, Netherlands Munchen, Germany Gainesville, FL, U.S.A.
R. Daams M. Hugueney P. Perez Gonzalez
Inst. de Geologia C.S.I.C. Universite Claude Bernard Universidad de Zaragoza
Madrid, Spain Lyon, France Zaragoza, Spain
M. Diaz-Molina L. Jacobs M. Petko
Universidad Complutense Southern Methodist Univ. University of Florida
Madrid, Spain Dallas, TX, U.S.A. Gainesville, FL, U.S.A.
W. Downs G. Koufos M. Pickford
Northern Arizona Univ. University of Thessaloniki Mus. Nat. Hist. Naturelle
Flagstaff, AZ, U.S.A. Thessaloniki, Greece Paris, France
629
z. Qui S.Sen I. Terlemez
Academia Sinica Universite P. et M. Curie M.T.A. Genel Mudurlugu
Beijing, China Paris, France Ankara, Turkey
M. Ringeade G. Storch H. Tobien
Universite de Bordeaux Forschungsinstitut Senck. Johannes Gutenberg Univ.
Bordeaux, France Frankfurt/Main, Germany Mainz, Germany
G. Sarac M. Sumengen D. Torre
M.T.A. Genel Mudurlugu M.T.A. Genel Mudurlugu Universita di Firenze
Ankara, Turkey Ankara, Turkey Firenze, Italy
R. Savage P. Tassy E. Unay
Bristol University Universite P. et M. Curie M.T .A. Enstitusu
Bristol, England Paris, France Ankara, Turkey
N. Schmidt-Kittler R . Tedford M. Woodburne
Johannes Gutenberg Univ. Amer. Mus. Nat. History University of California
Mainz, Germany New York, NY, U.S.A. Riverside, CA, U.S.A.
Participants of NATO ARW on European Neogene Mammal Chronology.
Front row (seated) left to right: V. Fahlbusch, M. Antunes, E. Unay, P. Mein, D.

Torre; second row (seated): R. Savage, A. Azzaroli, J. Morales, L. Ginsburg, E.
Martinez-Suarez, A. Agusti, N. Opdyke, T. Kotsakis, S. Moya-Sola, M. Pickford; third
row (standing): C. Guerin, S. Sen, A. van der Meulen, M. Alvarez-Sierra, M. Hugueney,
M. Freudenthal, C. de Giuli, J. Barry, G. Hock-Daxner, 0, Fejfar, F. Masini, M.
Bonifay; back row (standing): L. Jacobs, L. Flynn, G. Koufos, z. Qiu, K. Heissig, L.
Benda, P. Andrews, R. Daams, R. Ziegler, B. Engesser, K. Kowalski, F. Steininger, E.
Heizmann, G. Storch, R. Bernor, E. Lindsay. Participants not in picture: B. Alpagut,
H. de Bruijn, H. Lippolt, P. Tassy and G. Rossner (Secretary).
630
SUBJECT INDEX
Acheulian Paleolithic implements, Z16 Al Jadidah fauna, 597

Acid preparation, 61Z Alberca fauna, 509
Ad Dabtiyah, Z40, 595 Alcocer (AC), 149, 464, 471, 47Z
Adriatic Gulf, 351 Alcocer 3B, 150
Aerotrain (see Airborne Railway), 163 Alcoy, 399
Affalterbach, 188 Algeria, 59Z
Afghanistan, 401 Algerian fauna, Z38
Africa, 11, 133, 135, Z37, Z38, Z41, Z44, Aliveri, 66, 37Z, 393
Z49, Z53, Z58, Z61, 345, 357, Aliveri-Kymi, ZZ, Z8
369,371, 37~415,4ZZ, 4Z6, Aljezar B, 109
4Z9, 431, 456, 557, 573, 587, Almazan Basin, 51, 464
589, 596, 597 Almenara, 388
Africa mammal localities, 588 (map) Alpine Arch, Z58, Z60, Z61
African bioprovinces, 401 Alpine chronology, 483
African biostratigraphy, 436 Alpine glacial chronology, 1Z1, 1Z4
African dispersal event, Z38 Amana, 390
African mammal fauna, 587 Am berieu section, 398
African microfauna, 401 Ampudia section, 53
African middle Miocene fauna, 56Z Amuwusu, 544
African Miocene fauna, 589 Anaseur Formation, 499
African taxa, 367, 368, 558 Anatolia, 68, 71, 7Z, 444 (map), 515
African-Eurasian dispersal events, 371 Anatolian assemblage, 65
African-Eurasian mammal exchange, ZZ Anatolian assemblage zones, 67, 68
African-European dispersal, 357, 371 Anatolian succession, 61
African-European dispersal histories, Anchitherium fauna, Z39
43Z (map) Ancient Burdigalian taxa, 159
Afro-Arabian continental plates, Z41, Anjou, 157, 159, 168
557 Antarctic glaciation, 560
Afro-Eurasian land links, 589 Anwil, 85, 183, 187, 190, ZZ8
Age assignment, 3, 4 Apparent diversity, 579
Agenais region, 139 Apparent turnover, 579
Agenian land mammal age, 9, Z1, 73, Aquitaine Basin, 9, 139, 140 (map), 141,
157, 161, Z41 145, 146, 148, 150, 151, 153,
Agenian sites, 59Z Z37, Z38, Z45, Z57
Agenian stage, Z7, 51, 5Z Aquitanian parastratotype, 140, 154
Agenian/Orleanian boundary, 16 Aquitanian parastratotype of Carry-le-
Agreda, 358 Rouet, 14Z
Aillas, Z1, Z7, 140, 143, 144, 146, 147, Aquitanian stage, 18, Z1, 143, 178, Z55,
149, 150, 153, 154 Z59, 589
Airborne Railway, 157, 168 169, 171, Aquitanian stratotype, 139, 154
173 Aquitanian transgression, 143, 153
Akca yollen zone, Z4 Aquitanian/Burdigalian boundary, 16,
Akdag, 61 140
Akhalkalaki, 34 7 Aquitanian/Burdigalian stratotype, 153
Akkulaevo, 410 Arabia, Z59
Al Jadidah, 596 Arabian plate, 589
631
Arafara Sea, 43Z Astaracian land mammal age, 90, 2.41,
Aragon, 2.56 2.48, 2.49, 618, 62.5
Aragonian faunas, 394 Astaracian rodent fauna, 105
Aragonian of Poland, 384 Astaracian stage, ZZ
Aragonian stage, 4, 2.5, 51, 52., 57, Astaracian/Vallesian boundary, 16
357-359, 361-363, 365, 366, Astaracic faunal unit, Zl
368, 369, 401, 40Z, 497, 517, Ateca, 57
6ZO, 62.1 Ateca ill, 2.56
Aragonian stage/superstage, 618 Atlantic Basin, 557
Aragonian superstage, 618 Australia, 1Z, 43 Z
Aragonian/early Vallesian beds, 385 Autol (AU), 51, 149, 464, 471, 472.
Aragonian/early Vallesian sites, 386 Autochthonous phyletic speciation, 414
Aragonian/Vallesian boundary, 53, 55, Autochthonous sympatric speciation,
495, 497 414
Aragonian/Vallesian transition, 54 Autroche village, 162.
Arapli, 2.48 Auvilliers Castle, 162.
Araya, 375, 376 Avaray, 158, 164
Araya Basin, 383, 393 Avenida do Uruguay, 2.55, 2.56
Arcille, Z4, 36 Axios Valley, 32.1-32.3, 32.5, 32.6, 32.8,
Arctic environment, 2.06 331-336
Areias da Quinta do Bacalhau, 2.54
Areias do Vale de Chelas, 2.56 Baccinello, 34
Arenas del Ray, 367 Baccinello V3, 2.4
Arezzo, 350 Badener Tegel clays, 2.17
Argilas axuis de Xabregas, 2.5 7 Badenian stage, 19, zz, Z3, Zll, 2.16,
Argilas do forno do Tejolo, 2.56 2.17, zzz
Arid environment of Asia, 2.04 Bulimina-Bolivina zone, ZZ, Zll, 2.16,
Arikareean land mammal age, 535 ZZ7
Armantomys-Praearmantomys group, lagenid zone, Zll, 2.16, ZZZ, ZZ7
383 Badenian transgression, ZZ7
Arondelli, 2.32. Badenian/Kosovian transgression, 2.16
Arondellian subage, 2.1 7 Badenian/Moravian transgression, 2.16
Arquillo, 87, 361, 399 Badenian/Sarmatian boundary, 19
Arroyo de Val, 3 59 Babe fauna, 544
Artenay, 73, 157, 158, 162., 166, 168, Babe River, 544
2.3 7' 2.39, 2.42., 2.45, 2.46, 2.56 Bahean land mammal age, 2.74
Artenay fauna, 173 Baigneaux, 173, 2.37, 2.39
Artenay group, 169 Baigneaux-en-Beauce, 157, 158, 163,
Artiodactyl faunal groups 168
LG-I-m, 358 Balearic Islands, 390
LG-IV-VII, 360 Balizac, Zl, Z7, 139, 140, 144, 146, 150,
LG-vm-xm, 360 153, 154
LG-XIV, 361 Balizac section, 144
Arvicola cantiana zone, 104 Ballestar, 376, 383, 384, 395
Arvicola-Microtus superzone, 104, 110 Ballestar Formation, 376
Arvicola terrestris zone, 105 Balneario fauna, 511
Arvicolid radiation, 107 Baluchistan, 2.38, 2.39, Z4Z, 2.46
As Sarar, 2.40, 595 Banco Real, 2.56
Asia, 11, 2.58, 339, 340, 344, 357, 401, Banda Daud Shah, 574, 576
42.9, 548, 618 Baode, 309, 544
Asia Minor, 2.59, 576, 589, 597 Baode County, 52.7, 532.
Asia Minor mammal localities, 588 Baode fauna, 545, 546
(map) Baode (Shaanxi Province), 545
Asian-European mammal dispersal, 106 Baodean land mammal age, 2.74, 308
Asiatic bioprovinces, 401 Barstovian land mammal age, 2.54, 2.74,
Asiatic origins, 588 2.90, 2.99, 310, 536, 551
Asiatic taxa, 367 Basarakavak fauna, 2.3
Assemblage zone, 5, 67 Bashkiria, 407
Association zone, lZO Bauge-Savigne, 160
Astaracian assemblage, 2.17, ZZZ, 2.30 Bavarian Molasse, 181, 183, 190, zzz,
2.30
632
Bavarian Molasse Basin, 181, 190 Black Sea, 401, 426
Bayraktepe, 496-498 Blancan fauna, 339
Baza Formation, 380, 381 Blancan land mammal age, 134, 274,
Beauce limestone, 157, 165, 168 290, 339, 340, 604, 606
Beaugency-Tavers, 173, 246 Blancan Hagerman fauna, 339, 340
Beaugency-Tavers group, 173 Blesois, 157
Beaugency-Tavers Sand group, 171 Blois, 157
Beaugency-Tavers Sands, 164 Blois group, 173
Beaulieu, 21, 28 Blow plankton zonation, 255
Belchatow, 193-195, 202 Bohemian localities, 213
Belenyenice, 22, 28 Bohemian massive, 214, 216
Bellenberg, 183 Bohemian Tertiary basin, 211, 214, 222
Belometcheskaya, 248, 540 Bonanza, 213, 216, 227
Beni Mella!, 387,451, 597 Bordelais, 146
Berbesti Slavesti, 340 Boreal zone, 205
Beremend, 101, 110, 232, 405 Borsodia-Dolomys stage, 101
Bergasa (BG), 149, 464, 471, 472 Borsodia-Villanyia superzone, 99, 101,
Bering land bridge, 550 102, 110
Beteke, 410 Boselaphini bovids, 3 71
Betic massive, 258, 260 Bossee, 158, 166
Bezian, 239, 245 Botardo section, 384
Bezian fauna (Gers), 164 Bou Hanifia, 23, 31, 296, 496, 497, 499
Bierkeller, 179 Bou Hanifia Formation, 501
Bighorn Basin, 604 Bou Hanifia section, 502
Bigneaux-en-Beauce group, 171 Boulder horizon of Bavarian Molasse,
Biharian faunas, 214, 217 183
Biharian land mammal age, 94, 101, Boundaries of land mammal ages, 609
103, 106-108, 110, 128, 131, Bouzigues, 55, 150
135, 216 Brachyodus dispersal event, 415
Biharian zones, 104 Bridger Basin, 604
Biochron, 5, 6, 7, 128 Bridger fauna, 606
Biochronologic age, 551 Bridger Formation, 607
Biochronologic correlations, 264 Brockhorizont (boulder horizon) of
Biochronologic framework, 91, 618 Bavarian Molasse, 181, 183
Biochronologic units, 94, 133, 606 Brunhes Magnetic Chron, 480, 483, 484
Biochronology, 2, 3, 5, 7, 49, 91-93, 96, Brunhes/Matuyama magnetic boundary,
101, 112, 527' 574, 608, 613, 559
618, 622, 625 Briittelen 2, 177-179
Bioevents, 133 Bubenhausen, 188
Biogeographic aspects, 625 Bucheggberg, 177
Biogeographic distributional changes, Bugti, 70, 240, 242, 574
413 Bugti Beds, 241, 243, 245, 248, 372, 537
Biogeographic faunal provinces, 10, 12 Bugti fauna, 238-240, 371, 538, 548,
Biogeographic problems, 10 549, 551, 583
Biogeographic realms, 413, 415, 416 Bugti Hills, 237, 246
(map), 431 Bugti proboscidean datum, 243
Biostratigraphic correlation, 560 Bulong, 530
Biostratigraphic dating, 608 Buluk, 240, 242, 245, 248
Biostratigraphic event, 55 7, 564 Buiiol (BU), 359, 462, 464, 471
Biostratigraphic framework, 562 Burdigalian dispersal event, 238
Biostratigraphic interval zone, 561 Burdigalian fauna, 239, 592
Biostratigraphic unit, 125, 128 Burdigalian regression, 22
Biostratigraphic zonation, 20 Burdigalian stage, 18, 22, 153, 238, 253,
Biostratigraphical succession, 476 254, 256, 257, 259, 261, 323,
Biostratigraphy, 2, 3, 5-8, 61, 511, 580, 536, 589, 595
615, 625 Burdigalian transgression, 17 8, 256
Biotopes, 126 Burdigalian/Langhian boundary, 16
Biozonation, 67, 96, 119,616 Burdigalian/Langhian stages, 377
Biozone, 5, 6, 124, 125 Burgtonna 2, 105
Birosse, 245, 246 Burtepe Formation, 64
Bissingen, 55 Byelometcheskaya, 368, 372
633
C. Almirall (see also Can Almirall), 2.3, Casablanca, 361, 367, 375, 376, 384,
30 387-391, 401
Cabriel Basin, 10, 507, 508 (map), 509, Casablanca karstic complex, 379, 400
512. Cases de la Valenciana, 359
Cabriel section, 508, 510, 513 Castell de Barbera, 360, 366, 378, 383,
Calabrian stage, 615 384, 386, 394
Calah, 384 Castilla la Vieja, 2.97
Calamocha, 51, 52, 55 Catalonian basins, 375, 383
Calatayud-Daroca, 359, 365 Catalonian (Vallesian and Turolian), 62.5
Calatayud-Daroca Basin, 366, 375, Catalonium superstage, 618
383-386, 392., 396, 497 Caucasus, 2.04, 549
Calatayud-Teruel Basin, 51, 53, 615, Caunelles, 2.1, 2.7, 146, 153
62.0, 62.1 Celleneuve, 35
Calcaire blanc de l'Agenais, 139, 143 Cenozoic mammals of North America,
Calcaire blanc Formation, 140, 143 606, 608
Calcaire de Beauce Formation, 157 Central Massif, 157, 340, 346, 475
Calcaire gris de l'Agenais, 139, 144 Central Paratethys, 18, 2.0, 2.3
Calcaire gris Formation, 143, 153 <;erdanya Basin, 375, 376, 384, 390, 400
Calcareous nannoplankton, 2.56 Cervenf, 2.2.7
Calcarios com Chlamys scabriuscula de Cetina (CE), 149, 462., 464, 470
Musgueira, 2.56 Cetina de Aragdn, 51, 358, 384, 472.
Calcarios da M~gueira, 2.56 Cevizljik Formation, 61, 64
valdag Group, 61 Ceyssaguet, 480
Calta, 400 Chagny, 12.6,346,347
Camallera, 398 Chalkoutsi, 334
Can Almirall (see also C. Almirall), Chamtwara, 589
360,377-- Charmoille, 31 Z
Can Canals, 359 Chattian stage, 2.41, 595
Can Casablancas, 379 Chao Chia Cha (Qingyang County), 52.7
Can Jofressa, 379, 384 Cheb Basin, 2.11, 2.13, 2.14, ZZZ
Can Llobateres, 86, 108, 32.8, 330, 331, Chelas 1,2., 2.9
360, 366, 378, 383-387, 391, Chelas z, ZZ
~95, 396 Chetougou, 541
Can Marti Vell, 366, 377, 378, 384, 392., Chevilly, 157, 158, 2.39
393 Chi Chia Kou (Baode County), 52.7
Can Mas, 359 Chiana valley, 350
Can Missert, 360 Chianda, 589
Can Perellada, 379 Chillac, 2.4, 38, 343
Can Petit, 376, 383, 384, 395 Chilleurs, 173
Can Ponsic, 96, 360, 366, 378, 383, 386, Chilleurs-aux-Bois, 157, 158, 161, 168
395 Chilleurs-aux-Bois group, 169
Can Simeo, 383 China, 11, 136, 2.40, 331, 345, 454
Can Vilella, 384, 390, 400 Chinese Cenozoic stratigraphy, 52.8
Canada del Castano, 384 Chinese mammal localities, 534 (map)
Canales (pond), 4 70 Chinese-European faunal correlations,
~andir, 2.3, 30, 368, 447 530
Canis etruscus dispersal event, 134 Chinji, 2.39, 559, 561
Caproides dispersal event, 90 Chinji Formation, 2.46, 558, 574
Caravaca, 387 Chios, zz, 2.9, 2.48, 451
Carbon isotope shift, 561 Chita Parwala-Gabhir section, 2.40
Carpathian foredeep, 2.03 Chitenay, 158, 161, 168, 173, 2.39
Carretil (CAR), 464 Chitenay-Les Beilleaux, 169
Carriere Cluzel, 149 Chitenay-Les Beilleaux group, 168
Carry-le-Rouet (Formation Chlum, 110, 2.13, 2.14
biodetritique de Sausset, Choerolophodont proboscideans, 2.49
niveau), 140 Chomutov, 2.13, 2.14, 2.18, 2.19, 2.2.1
Casa del Acero, 2.3, 33, 360, 380, 384, Chorora Formation, 502.
385, 398 Chou Kou Tien, 52.7-52.9
Casa del Rincon (Rincon 1), 2.4, 36 Chron 5, 2.4, 1 2.8
Casa. Frat a, 346 Chron 5 (C3A), 2.0
Chron 9, 17, 19
634
Chron 11, 16, 17, 19 Cretaceous North American land
Chron 11/10 boundary, 19 mammal ages, 606
Chron 22r, 18 Crete, 502
Chron C3A, 19 Creux de Peyrolles, 346
Chron C5A, 19 Crevillente, 1-3, 23, 32, 360, 375, 384,
Chron C5Cn, 19 391, 398
Chron C5N, 16, 274 Crevillente 2, 87
Chron C5r, 23 Crevillente 6, 24, 33
Chron C6Br, 18 Cricetid vacuum, 52, 53, 55, 56, 71, 358
Chronologie framework, 255, 605, 612, Cricetodon-Hispanomys fauna, 385,
615 395, 497
importance of, 4 Cromerian glacial stage, 105, 107, 405
Chronostratigraphic dating, 587 Crostolo River, 347
Chronostratigraphic framework, 559 Csarn6ta, 110, 232, 405, 407, 409
Chronostratigraphic mammal stages, 25 Csarn6ta 2, 100, 109, 110
Chronostratigraphic stage, 24, 608 Csarnotan land mammal age, 101
Chronostratigraphic unit, 24, 25, 55, Csarnotan stage, 615
125, 128, 562 Ctenodactyloids, 69
Chronostratigraphy, 5, 6, 24, 92 Ctineves, 211, 214, 232
Chronozones, 5, 7, 8, 124 Cullar de Baza, 349, 400
Cixian, 529, 530, 540, 542 Czech karst, 214
Cladistics, 627 Czechoslovakia Neogene sites, 213
Clarendonian land mammal age, 274, (map)
296, 302, 310
Clark Fork Basin, 606 Dachai fauna, 547
Clarkforkian land mammal age, 7, 606 Dacian/Romanian boundary, 20
Climatic crisis, 483 Dalai Nor, 529
Climatic events, 5 Dam Formation, 595-597
Climatic fluctuations, 480, 482 Dama nestii dispersal event, 135
Climatozones, 124 Datum planes, 613
Cocumont, 140, 144, 146-150, 153, 154 Daud Khel, 574
Coderet, 146 De Meeren, 37
Coderet-couche, 154 Deep sea hiatuses, 560
Coelodonta antiquitatis-Bos primigenius Deinotherium dispersal event, 260
fauna, 529 Deinotherium reimmigration, 188
Coenozones, 120, 124 Dendil, 70, 516
Colhuehuapian land mammal age, 12 Deneze-sous-le-Lude, 158, 159
Collet Redon, 190 Dental morphology (see also Teeth
Comanesti 1, 23, 30 characters), 110, 111, 153,
Comanesti 2, 31 161, 239
Concud, 360 A/L indices of Vander Meulen, 103
Concurrent range zones, 6 bilophodont, 244
Condom, 239 brachyodont, 328, 331
Conohyus dispersal, 359 cementum, 231
Continental chronostratigraphic stages, enamel synclines, 231
20 height of crown, 284
Continental glaciations, 475,480 hypsodonty, 204,328,331,348
Continental plate movement, 557 mandibular incisor, 286
Continental/marine correlation chart, mesodont-lophodont molars, 107
16 rooted molars, 204
Continental/marine correlations, 21 rooted and rootless molars, 232
<;::orcoles, 359 supernumerary teeth, 243
Correlation group, 16 trilophodonty, 244-246
Correlation point, 153 Deseadan land mammal age, 12
Correlation tables, 57 Detan, 213, 214
Correlation tie-points, 26-31, 33-38 DeviD.ska Nova Ves, 22, 29, 211, 213,
Costablanca, 377, 381, 391, 392 216, 217, 220, 222, 223-229
Cournon, 536 Dhok Pathan Formation, 558
Cracow, 193 Diachronous, 3-5, 10, 53, 71, 123, 126,
Cremohipparion lineages, 315 240, 248, 254,497, 579, 612
Diavata, 323, 325, 332
635
Dinar-Akcakoy, 2.4, 34 Elephantoid datum event, 2.37
Dinarid-Asian giant rhinoceros region, Elmali Dag thrust, 61, 64
549 Elsterian sub age, 105
Dinotheriensande, 2.46 Empoli, 409
Dispersal events, 71, 12.6, 414, 549, 567, Emporda, 375, 376
602., 609 Emporda Basin, 398
Dispersal from Africa, 351 En Pejouan, 2.46
Dispersal from Asia, 351 End-Villafranchian event, 346
Djilancik, 2.39 Endemic Asian taxa, 567
Djilancik Beds, 2.46 Endemic fauna, 549
Dolinskoe, 410 Endemic taxa, 126
Dolna Cave, 2.04, 2.05 Endemism, 12.3
Dolnice, 2.13, 2.16, 2.19, 2.2.1-2.2.6 Ephemeral oxbow lake habitat, 560
Dolnice 1, 55 Eppelsheim, 245, 2.46, 312., 328, 330,
Dolnice 3, 2.11, 2.14, 2.18 396
Dongxiang Autonomous County, 537 Equid facial morphology
Dongyaozitou, 547 buccinator fossa, 282
Dor Zaggut sections, 592. canine fossae, 282.
Dordogne prehistoric sites, 482. caninus fossa, 283
Dorn-Diirkheim, 2.30, 396 fossette ornamentation, 284
Dou~-la-Fontaine, 166 infraorbital foramen, 2.82
Doupov Hills, 2.14 lacrimal, Z7 9
Douvre, 366 malar fossa, 283
Dragonian land mammal age, 606 nasal notch, 283
Dryopithecine hominoids, 96 orbital surface of lacrimal bone, 2.79
Duero Basin, 53,357,375,383,387,396 preorbital bar (POB), 279
Dumlupinar, 2.2., 2.9 preorbital fossa, 280, 2.83
Dverce, 2.13, 2.14 Erkertshofen, 55, 386
Dytiko (DTK), 32.1-32.3, 325, 328, 32.9, Er langgang, 543
334 Ertemte (Nei Mongol), 109, 527, 52.9,
Dytiko Formation, 323, 331, 332, 336 530
Dzerava skala, 110 Ertemte fauna, 528, 546
Dzungar Basin (Anjihai), 535, 536, 543 Escobosa, 360, 383, 385, 394
Eskihisar, 545
East Africa, 135, 239, 2.40, 245, 248, Eskihisar pollen assemblage, 66
413, 42.3, 42.8, 433, 435, 438, Eskihisar pollen zone, 2.2., 23
502, 589, 592-594 Esme-Akcakoy, 23, 31, 335, 369, 496,
East Africa Faunal Sets I and II, 590 497, 545
East African fauna, 592., 594 Estrepouy, 21, 2.8, 55, 140, 146, 150,
East Asia, 260 152., 153, 239, 2.56
East Paratethys, 2.39 Estrepouy (Gers), 144
Eastern Mediterranean, 20, 248 Estrepouy zone, 239
Ebershausen, 188 Esvres syncline, 157, 160, 161
Ebro Basin, 51, 359, 383, 462, 464 Etampes limestone, 157
Eburonian pollen zones, 136 Ethiopean biogeographic realm, 416,
Eburonian stage, 103 417, 42.0, 429, 43o-432, 435
Ecological factors, 10 Etouaires, 24, 36, 125, 12.6, 475
Eemian/Weichselian boundary, 105 Etulia, 410
Egerci Formation, 64 Eucladoceros dicranios dispersal event,
Egerian stage, 548 135
Eggenburg, 27 Eucricetodon gerandianus-
Eggenburgian stage, 18, 21, 2.14, 218, infralactorensis lineage, 146
548, 549 Eucricetodon haslachensis lineage, 146
Egypt, 589, 592, 594 Eurasia, 557
Eibiswald, 22., 2.8, 246 European land mammal age, 603
Eichkogel, 108, 109, 230 European mammal chronology, 615
El Canyet, 359,377,393 European mammal stages, 603
El Firal, 376 European marine stages, 609
El Kohol, 238 Eustatic sea level oscillation, 560
Elbistan, 24, 35 Euzet, 613
Elephant-Equus dispersal event, 341 Event stratigraphy, 4
636
Evolutionary change, 47 Freshwater Molasse (continued)
Evolutionary first occurrences, 12.8 Altere Serie, 181, 190
Evolutionary grade, 3 Hangendserie, 181, 183
Evolutionary lineages, 12.0 Jiingere Serie, 181, 190
Evolutionary processes, 12.6 Mittlere Serie, 181, 183, 188, 190
Extinction, 579 Freshwater Molasse of Bavaria, 182.
Extinction events, 561, 567 (map)
Fuenmayor (FM), 464
Facial morphology Fuente del Viso, 509, 512., 513
buccinator fossa, 342. Fuente Podrida, 512.
Falun, 158-160, 168, 173 Fuente Podrida fauna, 511
Falun of Anjou, 160, 165, 168, 171, 173 Fuente Podrida Formation, 510
Falun of Area Fugu County, 52.7, 545
Falun of Pontlevoy, 160
Falun of Touraine, 168, 171 Gaiselberg, 2.3, 31
Falun pit, 165 Gaiselberg land mammal age, 2.96
Falun transgression, 160, 166 Gaj Formation, 558
Fangshan, 530, 539 Gaj section, 574
Fangxian, 543, 549 Galerian faunas, 3 50
Farneta, 356 Galerian immigrants, 347, 349
Farneta fauna, 356 Galerian interval, 348, 349, 351, 356
Fars Formation, 589 Gallenbach, 188
Faunal distribution, 12.1 Gallenbach sand quarries, 183
Faunal events, 562. Gans, 2.1, 2.7, 140, 143, 146, 147, 149,
Faunal interchange, 4 150, 153, 154
Faunal provincialism, 12.3 Gansu Province, 52.7, 52.9, 532., 537,
Faunal succession, 53, 56, 57, 67 540, 549
Faunal zones, 561 Gaozhuangfauna, 546
Faunizones, 614 Gaozhuangian land mammal age, 2.74,
Fayum, 2.38, 589 308
Fayum Depression, 2.38 Garkin, 2.3, 32.
Fayumian land mammal age, 11 Gatinais limestone, 157
First Appearance Datum (FAD), 2.0, 94, Gauss/Matuyama magnetic boundary,
609 476
Fission track age determinations, 559 Gebel Zelten, 2.46, 368, 372., 592.-594,
Fissure fillings, 49, 55 597
Flood basin deposits, 462. Gebel Zelten fauna, 594
Fluvial channels, 462. Geiselberg, 2.30
Forest Bed series, 347, 348 Gemerek, zz, 2.8, 61, 63, 66, 70, 517,
Formation bioc1astique de Carry-nineau 518
13, 154 Gemerek area, 516 (map)
Formation biodetritique de Sausset- Gemerek mammal fauna, 52.4
niveau 2.1, 154 Gemerek section, 519-52.4
Formation pararecifale du Cap de Geochronologic age, 2.4, 608
N autes-niveau 7, 154 Geochronology, Z
Fornace RDB, 2.4 Geographic limits of land mammal
Forsthart, 2.2., 2.2.2. ages, 602.
Fort Ternan, 451, 597 Georgia (USSR), 345, 347
Fortuna Basin, 360, 367, 375, 376, 379, Gers, 2.46
398,400 Gier Bu Hasciso Formation, 592.
Four a Chaux quarry, 165 Giggenhausen, 2.14
Frankfurt Nord Basin, 2.56 Gilbert magnetic chron, 18
Frantiskovy Lazne (Franzensbad), 2.13 Glenns Ferry Formation (Idaho), 340
Franzensbad, 2.11, 2.14, 2.18, ZZO, Gliwice, 193
2.2.2.-2.2.9 Global climate events, 560
Franzensbad fauna, 2.11 Globigerinoides, 19, ZZ
Freshwater Molasse (see also Lower Globigerinoides lineages, 139
Freshwater Molasse, Upper Globorotalia aemiliana zone, 2.4
Freshwater Molasse, and Globorotalia inflata zone, 2.4
Siisswaserschichten), 181, 183, Globorotalia praemenardi-Globorotalia
188, 2.2.2. peripheroronda, 19
637
Globorotalia puncticulata/Gr. aemiliana Heterochrony, 608
transition, Z4 Heterogeneity of faunas, 1Z6
Goldberg, Z3, 30 Hiatuses, 6
Gombasek, Z13, Z17 High-resolution chronology, 10
Gomphotherium datum event, Z60 Himalayan foothills, 345
Gomphotherium dispersal event, 4 Himalayas, 557
Gomphotherium-Sayimys-Kansupithecus Hintersteinbruch, 179
level, 5Z9 Hinzirdag, 61
Gonen Basin, 445 Hipparion biozones, 336
Gonghe Basin {China), 136 Hipparion datum, 9, 10, Z5, Z74, Z9Z,
Gorafe, 109, 387 415,435,495,497,498,500,
Gorafe-Hue1ago Formation, 380, 381 50Z, 574
Gotzendorf, Z30 Hipparion dispersal event, 1Z6, 364,
Granada Basin, 367 495, 496
Granada depression, 349 Hipparion facial morphology
Grand Combe, 476 orbit-preorbital fossa, 330
Grande Coupure, 588 Hipparion fauna, 530, 541, 545
Grande Pile pollen sequence, 48Z Hipparion morphological terms, Z75
Grosslappen-Aumeister, ZZ8 Hipparion red clay, 5Z9
Great American Interchange, 5 Hipparionine, Z64, Z70, Z78, Z89-Z9Z,
Gres de Bazas, 144 Z94, Z98, 303, 306, 309-311,
Group 1 hipparionines, Z75, Z9Z, Z93, 314
Z96, Z99, 30Z, 307, 312-315 Hipparionine characteristics, Z75
Guadix Formation, 380 Hipparionine horses, Z63
Guadix-Baza Basin, 375, 376, 380, 384, Hipparionine lineages, 315
387, 391, 400 Hiramaki Formation, Z39, Z46
Guangxi Zhuang Autonomous Region, Hirschthal, 178, 179
5Z7 Hispanotherium fauna, Z55-Z57, 359
Guizhong, 530 Hiwegi, Z45, 589
Gulf of Loches, 166 Hofuf fauna, 597
Giilyazi, Z4, 36 Hofuf Formation, 451, 595, 596
Gumba, 589 Holsteinian sub age, 105
Gundersheim, 110 Hol;tejn, Z16
Gundersheim fauna, 406 Horlak, 516
Gundersheim-Findling, 405, 406, 408, Horlak 1, 66
409-411 Horlak Z, 64
Gundersheim-Findling fauna, 406 Hornomoravsky uval, Z16
Horta das Tripas, Z54, Z56
Hadrukh fauna, 597 Horta das Tripas fauna, Z55
Hadrukh Formation, 595 . Hostalets, Z96, 385, 386
Hagerman quarry, 339 Hostalets de Pierola, 360, 378, 383, 496
Hajna&.a, 10Z, 110, Z11, Z13, Z17 Hovorany, Z3, 31, Zl3, Z16
Halamagai Formation, 535, 543 Howenegg, Z3, 31, Z78, Z94-Z96, 310,
Halmyropotamos {Evia), 334 31Z, 313, 315, 396, 496, 497,
Hammerschmiede, 108, Z30, 396 499
Hannoba basalt, 536 Hrabrsko, 335
Hans Johnson Quarry, Z94 Hsanda-Gol, 53Z, 535
Harami, 61, 64, 65, 7Z Hsi-shui, 538
Harami 1, 66 Hsin Chia Kou {Quingyang County), 5Z7
Harami 3, 66, 67 Huai River, 548, 549
Hautimagne, 409 Hubei Province, 543
Headonian stage, 49 Huerta Obispalia {HO), 464
Hebei Province, 5Z9, 536, 540
Hefeng {Jingle County), 547 Iberia, Z55, Z58, Z60, Z61
Hemingfordian land mammal age, Z74, Iberian climate, Z60
Z90 Iberian fauna, 398
Hemphillian land mammal age, 134, Iberian Peninsula, 173, Z59, 350, 375,
Z74, 339 376, 384-386, 39Z, 395, 399,
Henan Province, 5Z7 401, 497' 550, 589
Hequ County, 5Z7 lbericus zone, 391
Hesperic massive, Z58, Z60 Ibero-Gentral Basin, 385, 387, 398
638
Ibero-Gentral province, 375, 395 Jhil Equus skull, 345
!hero-Levant fauna, 393 Jiangning County, 539
!hero-Levant province, 375, 377, 383, Jiangsu Province, 530, 538, 544
384 Jiaozigou, 537
!hero-Levant subprovince, 387, 392, Jingle, 529
394-397' 400, 401 Jiulongkou, 541
Ibero-Qccitanian province, 391, 399, Jiulongkou fauna, 540, 541
401 Jozaria, 238
Igdeli, 64, 70, 516, 517 Jurassic limestone, 205
Ikebulage Formation, 532
Immigrant-derived taxa, 567 Kabarsero section, 502
Immigrant taxa, 128 Kadzielnia, 194, 202, 206, 232
Immigration (see Dispersal) Kaiyuan Province, 530, 544, 549
Immigration from Africa, 351 Kalekoy, 64, 66, 516
Immigration from Asia, 351 Kalekoy assemblage, 70
Index fossils, 94 Kalekoy mammal fauna, 524
India, 416, 434, 454, 455, 457, 557, 560 Kalekoy section, 517-521, 523
Indian Basin, 557 Kalimantsi, 335
Indian Ocean, 589 Kamlial Formation, 239, 240, 558, 561
Indo-Pacific plate, 557 Kamyk, 194, 202, 206, 207
Indo-Pakistan subcontinent, 258, 259, Karaburun, 24, 35
496, 500, 549, 550, 557 Karaganian interval, 239
Indonesian Archipelago, 43 2 KaraozU, 66, 516
Indus River, 559 Karaozii assemblage, 70
International Code of Zoological Karaozii mammal fauna, 524
Nomenclature, 128 Karaozii section, 517-521,523
International Stratigraphic Guide, 5-7 Karpatian fore-deep, 216, 222
Inzersdorf (see also Vosendorf), 23, 31 Karpatian stage, 18, 19, 22, 214, 216,
Ipolytarnoc;-239-- 222
Ipolytarnoc fauna, 239 Karstic sites, 216
Iran, 589 Karungu, 58 9
Irvingtonian deposits, 350 Kastellios, 23, 31, 334, 335, 496, 497
Irvingtonian land mammal age, 604, 606 Kastellios section, 502
Ischim-Petropavlovsk, 109 Kayadibi, 23, 32, 335, 369
Isochronous, 6, 8 Kayseri-Sivas Basin, 61, 62, 515
Isochronous boundary, 121 Kazakhstan, 239, 246
Isochronous surface, 120 Kenya, 11, 417, 587-589, 592, 595, 597
Isochrons, 4 Kenyan fauna, 597
Isochrony of chronostratigraphic zones, Keypar, 303
562 Khapry, 341
Isotopic chronology, 264 Kiahera, 589
Isotopic dating (see also Radiometric Kielniki, 194, 202, 206, 207
dating)-;-z,-s- King-Yan-fou (China), 71, 390
Isotopic events, 560 Kinik, 23, 33
Isotopic excursions, 561 Kiramaki Formation, 242
Israel, 587, 594 Kirthar Range, 557
Itaboraian land mammal age, 12 Kiscelian stage, 214
Ivanane, 335 Kishinev-Odessa region, 410
Ivanovce, 110,211,213,217,224,225, Kislangian land mammal age, 101
231, 232, 407' 409 Kizilhisar pollen zone, 23
Ivanovce A, 109, 110 Kiziltepe metamorphic series, 445
Ivanovce B, 110 Kohfidisch, 90, 204, 230
Kohat-Potwar region, 558
Jabal Midra ash-Shamali, 595 Kolfnany, 110, 211, 213, 217, 232
Japan, 240, 258, 260, 429, 430 Kopeprusy, 110, 213, 214
Jaramillo magnetic subchron, 345, Koru, 240, 589, 592
347-349 Kosovian assemblage, 217
Javoricko, 216 Kotlovina, 410
Jebel el Quatrani Formation, 589 Koujiacun fauna, 542
Jebel Qatrani fauna, 238 Koujiacun Formation, 540
Jettingen, 183 Krakow-Wielun Upland, 203-205
639
Krolewskie, 2.02. Lantian Formation, 544
Kromdraai, 135 Lanzhou, 551
Kromdraai A, 133 Lanzhou Basin, 537
Kromidovo, 335 Lanzhou fauna, 53Z, 535
Kryzhanovka,410 Lanzhou railway station, 532.
Kiid,tk-Cekmece, 3Z Las Planas, 365
Kulu, 2.45 Last Appearance Datum (LAD), ZO, 94
Kurosedani (Japan), 42.6 Laugnac, 2.7, 82., 139, 140, 144, 146,
Kuruksaf, 344 150, 153, 154, 169, 536
Kyjov lignite seam, Z16 Le Bardon, 164
Le Coupet, 133, 136
La Alberca, Z4, 33, 361, 391 Le Coupet fauna, 547
La Aldehuela, 399 Le Moune, Z45
La Bastida, 366, 376 Legetet, Z40, 589, 592.
La Brete, Z7, 140, 144, 146, 152.-154 Lengshuigou fauna, 541, 542.
LaBrosse, 159, 160 Lengshuigou Formation, 530
La Buste, 359 Leptobos etruscus dispersal event, 136
La Canera, 360 Les Beilleaux, 159, 160, 168, 173
La Celia (see also Los Gargantones), 2.3, Les Cevennes, 153
3Z, 379, 391, 398 Les Echets pollen sequence, 482.
La Celia Basin, 360 Les Etouaires (see Etouaires)
La Chaux, 149, 150 Les Forques, 55--
La Ciesma, 359 Les Grandes Noues, 165
La Dehesa, 53 Lespignan, 2.7, 153
La Fortesa, 379 Levkon, 2.3
La Galocha (GAL), 464 Levkon 1, 32.
La Grenatiere, Z3, 30 Librilla, 33, 361, 384, 511
La Grive, 190, 394 Libya, 592., 594
La Grive M, Z3, 30 Libyan fauna, 597
La Hornera, 34, 380 Lineage zone, 1 ZO
LaMilloque, 140,146,150,154 Linqu County, 539
La Moliere, 177 Linxia Basin, 537
La Paillade, 153 Lisboa, Z38, 2.39
La Puebla de Valverde, 343 Lisboa IVb, ZZ, 2.8
La Romieu, 83, Z39, Z45, Z56, 6Z1 Lisboa Va, ZZ, 2.8
La Tour, 109 Lis boa Univ. Catolica, Zl, Z7, 153, Z55,
Lachar, 349 Z56
Lagenid zone, ZZ Lisbon Basin, 173, 2.54
Lagrelius Collection, 5Z8, 5Z9 Lisbon series, Z55-Z57
Laimering, 188, 190 Lisov, 2.13, 2.14, ZZ6
Laimering clay quarries, 183 Lithofagous pholads and lithodomes,
Lake Baykal, 407 165
Lake deposits, 46Z Lithostratigraphic framework, 559
Lakhuti fauna, 348, 350 Lithostratigraphy, 61
Laki Range, 557 Livenzovka, 341, 410
Lance Cretaceous interval, 606 Livenzovka Equus skull, 34Z
Land mammal ages, Z, 5, 7, Z1 Local fauna, 1Z7
Langebaanian land mammal age, 11 Loire Basin, 9, 157, 158, 168, 170 (map),
Langebaanweg, 134 172. (map) 173, 2.37, 2.38, 2.45,
Langenmoosen, 183, ZZZ, ZZ4-ZZ7 2.54
Langhian falun of Touraine and Anjou, Loire River, 158, 162., 164
166 Loperot, 2.48
Langhian part of the falun, 15 7 Loranca Basin, 358, 462., 464, 470
Langhian regression, ZZ Loranca del Campo, 358
Langhian stage, 19, 158, 2.54, 2.59 Los Gargantones (see also La Celia), 2.3,
Langhian transgression, 385 3Z, 360 - - -
Langhian/Badenian transgression, ZZ Los Mansuetos, 87, 98, 99, 109, 360, 385
Langhian/Serravallian transgression, Lothagamian land mammal age, 11
2.60, 2.61 Lower Freshwater Molasse (see also
Languedoc region, 153 Freshwater Molasse), 177, 178
Lantian area, 530, 544
640
Lower Freshwater Molasse reference Mammal Neogene (MN) zone 3, 82, 214,
faun~ 177 222, 597
Lower Miocene fauna, 55 Mammal Neogene (MN) zone 3~ 178
Lower Silesi~ 203 Mammal Neogene (MN) zone 4, 66, 83,
Lowest Stratigraphic Datum (LSD), 609 222, 237, 2'48
Luc sur Orbieu, 23, 30 Mammal Neogene (MN) zone 4~ 179
Lufeng (Yunnan Province), 530, 545 Mammal Neogene (MN) zone 5, 83, 173,
Lukavice, 213, 214 183, 190, 214, 248
Lumiar, 253 Mammal Neogene (MN) zone 6, 84, 222
Lunel Viel, 350, 356 Mammal Neogene (MN) zone 7, 84, 190,
Luoyang (Henan Province), 527 203
Lutetian transgression, 589 Mammal Neogene (MN) zone 8, 85
Mammal Neogene (MN) zone 9, 86, 214
Maboko, 240, 248, 368, 433, 592, 595, Mammal Neogene (MN) zone 10, 86
597 Mammal Neogene (MN) zone 11, 87
Maboko fauna, 593 Mammal Neogene (MN) zone 12, 87
Macedonia, 321, 333-335 Mammal Neogene (MN) zone 13, 87
Macromammal events, 124 Mammal Neogene (MN) zone 14, 88, 92
Maghreb, 416 Mammal Neogene (MN) zone 15, 88, 9Z
Magnetic Anomaly 3A, 24 Mammal Neogene (MN) zone 16, 89
Magnetic Anomaly 4, 18 Mammal Neogene (MN) zone 17, 89, 214
Magnetic Anomaly 5, 16, 17, 19, 23 Mammal Neogene (MN) zones 4-5, 214
Magnetic Anomaly SA, 19 Mammal Neogene (MN) zones ~6, 193
Magnetic Anomaly 5C, 19 Mammal Neogene (MN) zones 5-7, 202
Magnetic polarity time scale (MPTS), Mammal Neogene (MN) zones 8-13, 203
16, 18, 501, 507, 510, 562, 566 Mammal Neogene (MN) zones 1~17,
Magnetostratigraphic correlation, 559 214
Magnetostratigraphic data, 500-502 Mammal Neogene (MN) zones, 15b-17,
Magnetostratigraphic dating, 559 217
Magnetostratigraphy, 2, 8, 10, 25, 507, Mammal Neogene/Quaternary (MNQ)
511, 515, 521, 524, 558 concept, 124
Maigen, 21, 27 Mammal Neogene/Quaternary (MNQ)
Makapan 3, 133 system, 128
Makapanian land mammal age, 11 Mammal Neogene/Quaternary (MNQ)
Mala Cave, 204 zonation, 119, 123, 124, 127
Malakoy Formation, 61 Mammal Neogene/Quaternary (MNQ)
Mallorca, 71 zones, 1Z0-1 zz, 1 Z5
Malu\lteni, 100, 340 Mammal Paleogene (MP) reference
Mammal age, 121, 127, 128 levels, 48, 49
Mammal associations, 120 Mammal Paleogene (MP) zone 8, 49
Mammal biochronology, 25, 48 Mammal Paleogene (MP) zone 9, 49
Mammal biochrons, 621 Mammal Paleogene (MP) zone 16-18, 49
Mammal biozonation, 61Z Mammal Paleogene (MP) zone 21, 214
Mammal dispersal event, 2, 4, 5, 9, 133, Mammal Paleogene (MP) zone 25-30, 49
475, 515 Mammal stage, 5-7, 121
Mammal episodes, 21 Mammal substage, 121
Mammal extinction events, 475 Mammal unit, 21, 127, 1Z8
Mammal faunal zone, 20 Mammalian fossil assemblages (MFA),
Mammal localities in Poland, 194 (map), 461
202 (map) Mammalian paleofaunas, 48
Mammal Neogene (MN) biochrons, 8 Mammalian succession of Chin~ 533
Mammal Neogene (MN) chronology, 47 Manchar Formation, 68, 71, Z45, 558,
Mammal Neogene (MN) faunal units, 21 574, 575
Mammal Neogene (MN) faunal zones, 3, Manchones, 90, 359, 366
6, 8, 9, 20, 21, 57, 74 Manthelan, 158, 166
Mammal Neogene (MN) zonation, 57, Marada Formation, 59Z, 594
61, n, 73 Maragheh, 23, 3Z, Z97, 30Z, 307, 309,
Mammal Neogene (MN) zone 1, 81 314,335
Mammal Neogene (MN) zone 2a, 81 Marine biochronology, 16
Mammal Neogene (MN) zone 2b, 82 Marine biostratigraphy, 613
Marine chronologie framework, 20
641
Marine planktonic biozonation&, 18 Mimomys (Mimomys) pliocaenicus zone,
Marine planktonic scales, 16 10Z, 107
Marine transgression in plankton zone Mimomys (Mimomys) polonicus zone,
N6, 59Z 10Z
Marine transgressive events, Z57 Mimomys (Mimomys) savini zone, 104
Marine/continental biochronology, Z1 Mimomys (Mimomys) savini-Mimomys
Maritsa, 109, 400 (Cseria) pusillus zone, 104
a
Marnes Ostrea aginensis, 139, 144 Mindel-Riss interglacial age, 350
a
Marnes Unios du Bazadais, 139, 143, Mindel-Riss interglacial fauna, 1Z1
144 Mino, Z39, Z46
Marsolan, 55, Z37 Miocene chronostratigraphy, 18
Marti Vell, 383 Miogypsinidae lineage, 139
Martini zone of NN Zone 4, Z57 Mira, 375
Mas Genegals, 407 Mirabeau, 158, 166
Masada del Valle .z, 109 Moghara, Z48, 594, 597
Masia del Barbo, 86, 98, 330, 331, 335, Moheda (MOH), 464,471, 47Z
360,397 Moissac, 140
Maspino gravels, 350 Moissac I, 143, 146, 147, 150, 153, 154
Masquefa, 360 Molasse de l'Agenais, 140
Massendorf, ZZ7-ZZ9 Molasse de l'Armagnac, 144
Matassino, 344, 346 Molayan (Afghanistan), 71
Matassino local fauna, 345 Moldavia, 350
Matuyama magnetic chron, Z4, 480, 547 Molf Calopa, 55, 358,377, 381, 391, 39Z
Mediterranean, 561 Molina de Segura, Z3, Z4, 33, 34, 367,
Mediterranean area, 4ZO, 4Z5, 495, 500, 380, 391, 400
50Z, 503 Moncalvillo (MON), 149, 464, 471, 47Z
Mediterranean area faunal intervals, Mongolia, 110
615 Mongolian Gobi area, 5Z8
Mediterranean basins, 376 (map) Mont Dorian eruptions, 476
Mediterranean Sea, 548, 550 Montaigu, 81
Mediterranean (Tethyan) stage, 18 Montaigu-le-Blin, 154
Megacricetodon extinction event, 497 Montalban, 384
Meigne-le-Vicomte, 159 Monte Gargano, 11 Z
Menorca, 384 Monticino (Brisighella), 34
Merkur-North, Z13, Z18 Montopoli, 37, 34Z
Merychippine grade horses, 264, 289, Montopoli fauna, 476
30Z Montopoli faunal unit, 340
Messinian salinity crisis, Z03, 370, 399, Montopoli-type faunas, Z4
550 Montpellerian interval, 91, 9Z
Messinian stage, 19, Z4, 379 Montpellier, 1Z5
Messinian/Zanclean boundary, 16, 18, Montredon, 108, ZZ8, 335, 385
zo Montrejeau, Z46
Meswa, Z40, Z4Z, 59Z Moratilla, 358
Meswa Bridge, Z40, Z41, 589, 597 Moratines, 359
Meta-Carpathian arch, Z03 Moravian region, Z13, Z16
Mexico, 339 Moreda, Z3Z, 399
Mfwangano, Z4Z, Z45, Z46 Morocco, 59Z, 597
Micromammal dentitions, 6Z7 Morosovka, 410
Micromeryx immigration, 359 Moroto, Z40
Microtocricetus molassicus zone, 98 Mortilla, 53
Microtus-Mimomys Superzone, 103, 110 Moruorot, Z40, 589
Mid-Vallesian boundary, 357 Mosbach-Z, 104
Mid-Vallesian crisis, 363, 364, 366 Mosbachian land mammal age, 1 Z8
Middle Miocene transgression, Z61 Mounicot, 55
Mikulov, Z13, Z16 Mt. Blanco, 606
Mimomys (Mimomys) davakosi zone, Mt. Luberon, Z98-300, 30Z
100 Mt. Vully, 178
Mimomys (Mimomys) hajnackensis zone, Muroid rodent molar evolution, 107
10Z Murree Formation, 71, 386, 558, 574,
Mimomys (Mimomys) occitanus zone, 576
101, 106 Mutenice, Z13, Z16, ZZ3,ZZ6
642
Mwiti, 2.40, 2.42., 2.45, 2.46 Nihewan land mammal age, 274
Myomiminae, 381 Ningxia (Wuzhong County), 530, 545
Ningxia Hui Autonomous Region, 540
Nagri Formation, 502., 558 Nombrevilla, 108, 328, 330, 331, 360
Nan Sha Wa (Hequ County), 52.7 Norfolk, 34 7, 348
Nannoplankton biozonation (see also North Africa, 136, 248, 428, 499, 576,
Plankton zone), 18 577, 616
Nannoplankton zone N4-N5, 153 North African faunal sequence, 592
Nannoplankton zone NN1, 18, 2.1 North America, 11, 134, 253, 254, 258,
Nannoplankton zone NN3, 18, 2.2 260, 261, 339, 342, 344, 350,
Nannoplankton zone NN4, 22, 257 429, 431, 496, 548-550, 604,
Nannoplankton zone NN4 floras, 19 608, 621, 622
Nannoplankton zone NN4/NN5 North American Code of Stratigraphic
boundary, 19 Nomenclature, 5, 7, 606
Nannoplankton zone NN6, 22, 216 North American gomphotheres, 246
Nannoplankton zone NN6/NN7 North American land mammal age
boundary, 19 concept, 119, 128
Nannoplankton zone NNll (Chron CN9), North American land mammal ages, 3,
18 7, 11, 601-603, 608-610
Nannoplankton zones NN4-13, 255 North American life zones, 605
Nannoplankton zone NN 11 a/NN 11 b North American provincial land
boundary, 19 mammal ages, 606
Nannoplankton zone NN1lb/NN12 North American tetralophodonts, 246
boundary, 19 North American vertebrate chronology,
Napak, 589 607
Narbada River (India), 345 North China, 527, 545, 546, 548, 549
Nari Formation, 557 Novaya, 410
Natural Remanent Magnetization Numerical taxonomic methods, 278
(NRM), 511, 517
Navarrete del Rio, 358 Occam's razor, 587
Navere, 239 Occitanium superstage, 618
Navere 1, 55 Oceanic circulation, 561
Nea Mesimvria Formation, 323, 330, Oggenhof, 183, 188, 190
331, 336 Olchon Peninsula, 407
Nearctic biogeographic realm, 416 Old World tropics, 568
Negev, 594, 597 Olduvai beds, 135
Negev fauna, 240, 594 Olduvai Gorge, 306
Nei Mongol Autonomous Region, 546 Olduvai magnetic subchron, 24, 73, 136,
Neogene chronostratigraphy, 18 480, 547
Neogene dispersal phase 1, 249 Oligocene/Miocene boundary, 18, 153,
Neogene dispersal phase 2, 248 415, 436, 462, 464
Neogene monophyletic groups, 242 Oligocene/Miocene transition, 139, 150
Neotropical biogeographic realm, 416 Olivola, 133, 343-345, 356
Neudorf an der March, 443 Olivola fauna, 133, 135, 136
Neudorf clay pit, 213, 216 Olivola faunal unit, 131, 132, 346
Neudorf fissures, 227 Ombo, 368
Neudorf Sandberg, 23, 29, 211, 213, Omo Basin, 438, 439
216, 217' 447 Oncophora (Rzehakia) beds, 222
Neudorf Spalte, 213, 541 Opole, 193, 195, 202, 203
Neudorf Spalte 1-3 (see also Devinska Oppelian zone, 120
Nova Ves), 22, 29 Orce-Galera-Huescar, 380
Neudorf Spalte 3, 217 Orechov, 22, 28, 211, 213, 216, 219,
Neuville-aux-Bois, 157, 158, 161 221-224, 226
Neuville-aux-Bois group, 169 Oriental biogeographic realm, 416, 417,
Ngorora, 433 420, 429, 431
Ngorora Formation, 369, 502 Oriental region, 550
Ngororan land mammal age, 11 Or.leanais Sands, 157, 161, 239, 246
Nihewan, 532 Orleanian assemblages, 214, 218
Nihewan fauna, 133, 345, 528, 529, 547, Orleanian Basin, 157
548 Orleanian land mammal age, 21, 73,
Nihewan Formation, 547 158, 239, 241, 245, 2.49, 415,
429, 593, 595, 618, 625
643
Orleanian stage, 27-31 Peralejos, 108
Orleanian/Astaracian boundary, 16 Peralejos D, 108
Orleanic faunal unit, 21 Peridyromys prosper-brailloni group,
Orleans, 157, 158, 162, 173 153
Orrios, 232 Perpignan (see also Serrat d'en
Orrios 3, 35, 110 Vacquer), 35, 88,90, 134,476
Ostracoda, 256, 257 Perpignan-Roussillon, 125
Osztramos, 407 Perrier plateau volcanic
Ottnangian stage, 18, 22, 211, 214, 216, chronostratigraphy, 479
222, 549 Perrier-Rocca Neyra, 476
Oued Zra, 23, 32 Perroche quarry, 165
Oxygen isotope climatic stage, 481 Petersbuch 2, 55
Oxygen isotope curve, 423, 483, 484 Petit Camon, 55
Oxygen isotope records, 561 Peyrolles, 475, 476, 480
Pfaffenhofen, 188
PII pollen zone, 24 Phylogenetic changes, 413
Pill pollen zone, 24 Phylogenetic systematics, 627
PIV-PII pollen zones, 24 Phylozones, 120, 124
Pacific Basin, 557 Piacenzian stage, 16
Pakistan, 11, 71, 136, 507, 557, 560, Piedra Formation, 376
573, 574 (map), 575, 577, 595 Piera, 297, 360, 391
Palearctic biographic realm, 416, 417, Piera section, 398
429-431 Pinjor "stage", 340
Palearctic zoogeographic province, 548 Pikermi, 23, 33, 109, 302, 325, 332, 335
Paleoclimatology, 475 Pikermian mammalian stage, 128
Paleogene biozonation, 8 Pithiviers limestone, 157
Paleogene chronostratigraphy, 18 Plakia, 23, 30
Paleogene/Neogene boundary, 18 Plankton zone N 5, 18
Paleogeography, 588 Plankton zone N7, 22
Paleomagnetic data, 136 Plankton zone N8, 22
Paleomagnetic dating, 601, 608 Plankton zone N9/N10, 22, 23
Paleomagnetism, 128 Plankton zone NlO, 19
Pannonian assemblages, 227 Plankton zone N12, 23
Pannonian Basin, 615 Plankton zone N15, 16, 19
Pannonian stage, 214, 216, 293 Plankton zone N16, 23
Pannonian strata, 23 Plankton zone N17, 23
P annonian/Pontian stage boundary, 19 Plankton zone N 17/N 18 boundary, 19
Panska G6ra, 204 Plankton zone NN1, 154
Papiol, 359 Plankton zone NP25, 18, 154
Paracuellos, 359 Planktonic biostratigraphy, 613
Parahippine horses, 264, 289 Planktonic foraminifera, 256
Parapodemus gaudryi zone, 99 Planktonic foraminifera zonations, 18
Parapodemus lugduneneis zone, 98, 99 Plate tectonics, 429, 433
Paratethys, 203, 238, 532, 548, 550 Platybelodont proboscideans, 249
Paratethys area faunal intervals, 615 Platyrrhine primates, 422
Paratethys basin, 211, 222 Plesivec, 110, 211, 213, 217
Paratethys sea, 216, 260 Pliocene chronostratigraphy, 18
Pardines, 476 Pliocene/Pleistocene boundary, 73, 214,
Paris Basin, 601, 609, 613 547
Parr ales (PAR), 464 Pliohyaena brevirostris dispersal event,
P~alar, 22, 23, 29, 245, 368, 372, 133
443-446,451-454,493 Plio-Pleistocene of western Europe, 122
P~alar deposits, 450 Po Valley (Italy), 615
Pa~alar fauna, 447, 455-457 Podlesice, 88, 100, 109, 194, 195,
Pastonian pollen horizon, 346, 347 202-204, 390, 407
Patagonia, 422, 430, 435 Poggio Mirteto, 36
Pataniak, 387 Poitiers Gulf, 166
Paulhiac, 27, 81, 140, 143, 146, 149, Polish lowland, 203
150, 153, 154, 536 Polish mammalian successions, 202, 207
Pedregueras, 108,391 Pollen zonations, 20
Pentalophos (PNT), 332 Polyphyletic theories, 126
644
Pond deposits, 46Z Qinling Mountains, 549
Pont Boutard, 166 Quantougou, 540
Pontian of 'Eurasia, 5Z8 Quaternary, 10
Pontian stage, Z16, 3Z3, 615 Quaternary glacial chronologies, 1Z3
Pontian/Dacian boundary, ZO Quel (QE1), 464, 471, 47Z
Pontign~, 158 Quinta da Farinheira fauna, Z55
Pontlevoy (= Pont Levoy-Thenay Quinta da Noiva, Z45
(Falun)), 83, 158, 161, 165, Quinta das Pedreiras, Z45, Z53
168, 173, Z45, 6Z1 Quinta do Navigao, ZZ
Pontlevoy fauna, 165 Quinta do Narigao fauna, Z55
Pontlevoy sea, 166 Quinta do Pombeiro, ZZ, Z9
Pontlevoy-Thenay Sand, 165, 171 Quinta do Pombeiro fauna, Z55
Portugal, 55
Potwar biostratigraphy, 575, 577 Radiometric data, Z39, 499
Potwar faunal change, 567 Radiometric dating (see also Isotopic
Potwar Plateau, Z46, 497, 50Z, 558, dating), 5, Z14, 559, 589, 59Z,
57 5-578, 583 601
Potwar Plateau fossil plants, 560 Radiometric scale, 57
Potwar sequence, 559, 563 Ramapithecine hominoids, 96
Potwar Siwaliks, 573 Ramblian fauna, 55
Poznan Formation, Z03 Ramblian stage, Z5, 51-53,57,357,358,
Pozo (PZ), 464 361, 368, 371, 6ZO, 6Zl
Pre-Aragonian zone X, 51 Rancholabrean deposits, 350
Pre-Aragonian zone Y, 51 Rancholabrean land mammal age, 606
Pre-Aragonian zone Z, 51 Range zone, 5, 1ZO, 1 Z4
Prebeza, 451 Rauscherod, ZZ, ZZZ
Prezletice, 110, Z14 Ravin de Konikovo, 3Z1
Primitivus zone, 39Z Ravin de la Pluie (RPI), 3ZZ-3Z4,
Proboscidean datum, 9, 11, 73, Zll, Z53 333-335
Proboscidean dispersal event, 4, Z37, Ravin de Vatillik, 3Zl
Z48, Z49, Z53, 4Z5 Ravin des Zouaves (RZ), 3Z1-3Z4, 333,
Prochoma (PXM), 3Z3, 3Z5, 334 334
Progonomys hispanicus zone Rebielice Krolewskie, 10Z, 110, 194,
Progonomys-Rotundomys superzone, 96, 195, zoz, Z05, 407
98, 108 Red Cloud Formation, 350
Promimomys insuliferus zone, 100 Reference faunas of land mammal ages,
Promimomys-Mimomys-Arvicola 6ZZ
lineage, 91, 94, 110, 111 Reference localities, 8
Promimomys-Mimomys stage Reference sections, Z
(Csarnotan), 100 Reisensburg symposium, 6Z7
Promimomys moldavicus zone, 100 Rema Marmara, 34
Promimomys zone, 100 Rembach, ZZ, ZZZ, ZZ8
Przeworno, 194, 195, Z03 Restricted species of land mammal
Przeworno fauna, Z03 ages, 6ZZ
Pseudotheridomys group, 151 Rhodanomys lineage, 150
Pseudotheridomys lineage, 150 Rhodanomys-Ritteneria group, 146,
Pseudotheridomys-Ligerimys group, 150 148, 150, 6Z1
Ptolemais, 109 Rhodos, 37
Ptolemais 1, Z4, 35 Rhone Basin, 387
Ptolemais 3, 100 Rhone valley, 398
Puebla de Valverde, 135 Richevoltes, 55
Puenta de Vallecas, 359 Rickenbach, 154
Pul-e Charki, 390 Ries crater, 183, 190
Puttenhausen, 183 Ries lake, 188
Pyrenees, Z60, Z61 Ries impact, 181
Pyrenees Basin, 384 Rincon, 34Z
Pyrenees Range, 375, 401 Rincon 1 (see also Casa del Rincon), Z4
Rio Cabriel, 5~
Qaidam (Tsaidam) fauna, 5Z9, 544 Rfp basalt cone, Zl4
Qinghai Province, 5Z9, 530, 535, 541 Riparian forest inhabitants, 98
QingyangCounty, 5Z7 Ritteneria lineage, 150
645
Riu Ripoll, 383, 397 Saudi Arabia, 451, 589, 594-597
Roca Neyra, Z4, 37 Saulcet, 150, 153
Rodent partial range zone, 1Z4 Savigne-sur-Lathan, 158, 159, 166
Rodent superzones, 94, 96 Savigneau facies, 158
Rodolfian land mammal age, 11 Schaffhausen, 153
Rombach, 66 Schernfeld, Z3Z
Ronzon zone, 613 Schnaitheim, 55
Ro ~haupten, 183 Sehwan section, 574
Rotem Basin, Z40, 594 Selles-sur-cher, 157, 158, 161 1 169
Roussillon Basin, 407, 615 Selva Vecchia, 346
Roussillon fa\Dla, 5Z8 Selvella, 344 1 346
Rubielos de Mora, 57 Semiarid to arid environments, 433
Rudabanya, 98, 108, Z30 Seneze, 73, 12.5, 133, 135, 136, 343,
Rlmton, 346, 348 346,475
Rupelian stage, 59 Z Seneze fauna, 132. 1 547
Ruscinian assemblage, Z31 Serrat d'en Vacquer (see also
Rus.cinian cricetid rodents, 110 Perpignan), Z4, 35, 88, 99, 100,
Ruscinian land mammal age, 16, Z4 1 110, 615
9Q-9Z, 94, 99, 100, 106-109, Serravallian stage, 19, 2.54 1 Z59
11Z, 1Z7, 1Z8, 134, Z04, Z74, Serravallian/Tortoni an boundary, 16
380,387, 405,411, 615 Ses Fontanelles, 390
Ruscinian stage, Z5, 34, 35 Sete, Z4, 35, 101, Z3Z, 409, 615
Ruscinic faunal unit, Z1 Seu d'Urgell, 375, 376, 384
Rusinga, Z39, Z45 Seu d'Urgell Basin, 366
Rusingan land mammal age, 11 Seventh International Geological
Rzehakia (Oncophora) beds, Z11, Z16 Congress, 604
Seynes, 12.7
Sabadell, 613 Shaanxi Province, 52.7, 540, 541
Sahabi, 136 Shagou (Nihewan s.s.) fauna, 547
Saint-Gerand-le-Puy , 536 Shandong Province, 5Z9, 539
Saint Vallier, 89, 1Z5, 343, 345, 356, Shanwang, 54Q-54Z, 551
475 Shanwang fauna, 530, 539, 551
Saint Vallier fauna, 133 Shanwang (Shandong Province), 52.9
Sainte Catherine, 55 Shanwangia, 539
Sakaraulian, Z39 Shanxi Province, 5Z7, 52.9
Salento Peninsular, 351 Shawan Formation, 536
Salobrena, 367, 384, 387, 391, 400 Shig-chiang-tzi-ku, 53Z
Salt Range, 558, 559 Shouyang, 308,309
Saltina Z, Z4 Siberia, 407
Samos, Z3, 3Z, 33, Z78, 309, 31Z, 331, Sicily, 71
332., 335 Sihong fauna, 538, 551
San Giusto, 409 Sihong Province, 530
San Juan (SJ), 464 Silesia, 193 1 ZOZ
San Men Series, 52.8, 52.9 Simbugino, 407, 410
Sandelzhausen, 181, Z46 Sind, 2.45
Sankt Stefan, Z3, 30 Sino-American project, 532.
Sansan, 84, 90, 185, 190, ZZ4, ZZ5, ZZ7, Sivalhippus complex, Z76, 302., 306, 307,
Z45, 359, 368, 451 311-315
Sant Andreu de la Barca, 55, 377, 381, Siwa, 597
391, 392. Siwa fauna, 594
Sant Mamet, 359, 377, 393 Siwalik assemblage, 580
Sant Quirze, 360, 366, 378, 383, 384 Siwalik biostratigraphic interval-zones,
Santa Cilia (SC), 147-149, 464, 471, 47Z 56Z
Santacrucian land mammal age, 4ZZ Siwalik biostratigraphic zonation, 561
Santarem, Z3, 30 Siwalik chronology, 583
Santiga, 360 Siwalik climates, 560
Sardice, Z13 Siwalik cricetid radiation, 583
Sardinia, 71 Siwalik extinctions, 568
Sarmatian stage, Z3, Z16, 32.3 Siwalik fauna, 566, 567, 582.
Sarmatian/Pannonia n boundary, 19 Siwalik faunal change, 566
Sarrion, 110, 399 Siwalik mammals, 562., 567
646
Siwalik murid record, 573, 577, 580 Stratigraphic signals, 4
Siwalik section, 576 Stratigraphic superposition, 3
Siwalik sequence, 497, 557, 564, 575, Stratophenetic series, 576
577, 592. Subzone, 1Z1
Siwaliks (see also Pakistan), 4, 11, 136, Suchomasty, 108, 2.11, 2.13, 2.14, 2.2.4,
368,447,451, 549, 557-559, 2.2.5, 2.2.7' 2.30
563, 567 Suevia fauna, Z14
Slatina 1, 37 Suevres Collonges, 179
Slatina 2., 37 Sulaiman Range, 558
Slovakia, 2.13 Suosuoquan Formation, 535
Small mammal diversity, 583 Superage/superstage, 62.5
Soan Formation, 558 Superposition, 2., 6-8, 12., 47, 49
Soblay, 366, 398 Superzone, 1 2.8
Sof~a, 2.3, 30, 2.48 Siissbrackwassermolasse, 179
Solar energy capture, 42.0, 440 Siissenborn, 348
Solar energy fluctuations, 42.2. Sut3wasserschichten (see Freshwater
Sologne Sands, 157 Molasse beds), 2.2.2.
Songhor, 2.40, 372., 589, 592. Swamp-marsh deposits, 462.
South Africa, 42.8 Synchronous events, 2.54
South America, 12., 2.54, 2.58, 42.9, 431, Synchrony, 4, 5, 8, 53, 495, 52.4
435 Systematic chron numbering, 18
South Asia, 573, 577, 580, 583, 584
South Asian biogeographic province, Taben-buluk, 538
575 Taben-buluk area, 52.9, 535
South China, 52.7 Tadjikistan, 348,350
South China (Yunnan), 548 Tagus, 365, 359, 383
Southeast Asian mammals, 562. Tagus Basin, 2.55, 2.57, 2.59, 2.61, 366
Southeast Asian species, 567 Tandem dental evolution, 413
Southern Asia, 2.38, 2.41, 2.45, 2.48, 2.49, Tandem evolution, 439, 440
608 Taphonomy, 487
Southern Asia elephantoids, 2.39 environments of deposition, 462.
Southwest Asia, 416 fossil concentrations, 559
Soviet Asia, 134 post-predator modification, 492.
Spalte, 2.2.0, 2.2.3-2.2.9 predator modification, 448, 488, 491,
Spalte-3, 2.16 559
Spanish extinction/immigration events, predepositional evidence, 452.
369 preservation state, 469, 470
Spanish Miocene bioevents, 370 scat and pellet concentrations, 559
Speciation event, 561 taphonomic factors, 10, 2.40, 465
Species associations, 62.5 Tarazona, 359
Sphaeroidinellopsis zone, 2.4 Tasso fauna, 135, 136
Spilia 1, 2.4, 35 Tasso faunal unit, 346, 356
St. Esteve, 134 Tavers, 158, 168, 173, 2.45
St. Genies, 2.2. Taxonomic boundaries, 415
St. Gerard, 358 Taxonomic diversity, 42.0, 433
Stage of evolution, 6, 8, 9, 1 z, 49, 613, Taxonomic diversity changes, 418, 419,
618, 62.5, 62.6 42.2., 42.4-42.6, 434, 436, 437'
Stampian stage, 536 440
Standard zone, 12.0, 12.4 Taxonomic grade, 2.45
Stehlin quarry (Artenay), 162. Taxonomic identification, 94
Steinberg, 2.3, 30 Tectonic events, 5
Steinheim, 84, 190, 2.46, 2.48 Teeth characters (see also Dental
Steinheim Basin, 190 morphology), 3, 102., 107, 110,
Stenotherm foraminifera, 2.59 112., 152., 2.39, 2.43, 333, 334,
Stephanomys rambliensis zone, 98, 99 347
Steppe lemmings, 107 antecrochet, 536
Sterkfontein, 133 anterior ellipse, 152.
Strakonice, 2.13, 2.14, 2.2.0, ZZZ, 2.2.3, 2.2.6 anteroloph, 148, 149
Stranska skala, 2.16 anterolophid, 69, 147, 152.
Stranzendorf, 2.4, 37, 38 anterosinusid, 152.
Stranzendorf D, 2.4 bunodont ctenodactylid, 68
647
Teeth characters (continued) Teruel-Alfambra Basins, 387
cheek tooth plications, 275 Tethys, 430, 589, 592.
chevrons, 576 Tethys area faunal intervals, 615
crown height, 3, 536 Tethys Sea, 2.38
dental morphology, 66, 70, 71 Tethys seaway, 42.9, 557, 588
ectostylids, 288 Thaler biozones, 613, 614
first syncline, 152. Thaler mammal zones, 612.
hypocone, 32.9 Thenay (see also Pontlevoy-Theney),
hypoglyph, 2.8 5 165, 166, 173
hypolophid, 152 Thermal demagnetization, 518, 52.1
hypsodonty, 108 Thessaloniki, 32.5
linguaflexid, 330 Thrace, 71
lingual cingulum, 536 Thvera magnetic subchron, 18, ZO
lingual cusps, 576 Tianshan Mountains, 536
lingual hypoconal groove, 3 2.9 Tibetan expedition, 530
longitudinal ridge, 147, 148, 152., 153 Tibetan Plateau, 530, 557
lophodont semi-hypsodont, 68 Tien-chiang-tzi-ku, 538
medisinus, 536 Tienshan, 52.7
mesoloph, 147-150, 152. Tiffanian land mammal age, 606
mesolophid, 69, 147, 152. Tiglian C, 2.4, 38
metalophulid, 71 Tiglian land mammal age, 136
molar ectoflexid, 2.89 Time succession, 47
paracone, 149 Time-transgressive dispersal, 94
parastylid, 32.9 Tongxin, 541, 542.
pli caballin, 2.85, 32.9 Tongxin area, 530
pli caballinid, 2.88 Tongxin fauna, 540, 541
plication, 32.9 Toringian land mammal age, 104,
posterior ellipse, 152 106-108, 110, 380
posterior cingulum, 536 Tornanian land mammal age, 101
posterior metalophule, 146 Torrejonian land mammal age, 606
posteroloph, 147, 148 Torremormoj6n section, 53
posterolophid, 69, 152. Torrent de Febulines, 360
postfossette, 2.84, 536 Torrijos, 359
premolar ectoflexid, 2.87 Tortajada, 109
premolar linguaflexid, 2.88 Tortajada A, 99, 109
premolar metaconid, 2.87 Tortonian stage, 19, 32.3, 499
premolar metastylid, 2.87 Transgression/regression cycles, 483,
protoconal spur, 2.86 475
protocone, 2.85, 32.9 Trilogie agenaise, 139, 143
protoconid, 69 Trilophomys-Ruscinomys Superzone, 99,
protostylid, 2.88, 32.9 109
sinus, 152. Trimingham, 348
sinusid, 152., 153 Trinxera Nord Autopista, 360, 384
talonid, 535 Trinxera Sud Autopista, 378, 397
third synclinid, 153 Triversa (see also Villafranchian), 2.4,
transverse metalophule, 146 36, 476
trigonid, 535 Triversa faunal unit, 340
Tegelen (see also Tiglian C), 24 Tropical Africa, 439, 455
Tegelen fauna, 133, 136 Tropical African faunas, 454
Teilchron, 5 Tsaidam (see Qaidam)
Teilhard de Chardin's Zone ll, 546 Tucho~ce, 55,Z13,Z14,Z18,Z19
Teilzone, 5 Tung-gur, 52.8, 52.9, 532., 540, 542.-544,
Teiritzberg, zz, 28 551
Temporal gaps, 6 Tung-gur Formation, 530
Ternanian land mammal age, 11 Tunisia, 592.
Terranova, 346 Tunisian fauna, 2.38
Terrassa, 379, 397 Turgai Strait, 548, 588
Terrats, 24, 34 Turkey, 71, 72., 2.48, 369, 454, 456, 616
Teruel, 613 Turkish plate, 589
Teruel Basin, 358, 360, 379, 397 Turkish-German lignite, 445
Teruel-Alfambra area, 615, 616 Turkomys-Byzantinia fauna, 395
648
Turolian assemblage, 513 Vallesic faunal unit, Zl
Turolian fauna, 360, 401 Valquemado (VQ), 358,464
Turolian faunal association, 396 Valtorres, Z56
Turolian dispersal event, 310 Vathylakkos (VLO), 3Zl, 3ZZ, 3Z4, 3Z5,
Turolian land mammal age, Z3, 90, 94, 3Z8, 334
98, 105-llO, llZ, Z30, Z74, Vathylakkos Formation, 3Z3, 331, 33Z,
Z97, Z98, 300, 308, 3Z9-33Z, 335, 336
336, 357, 361, 36Z, 364, 367, Vcelare, Zll, Zl3, Zl7, Z3Z
370,37Z,384, 389,398,399, Ycefare 3, 110
40Z, 407, 508, 615, 6Z5 Vendargues, Z4, 35
Turolian localities, 400 Venta del Moro, Z3, 33, 361, 384, 391,
Turolian stage, Z5, 3Z-34, 134 509, 5ll .
Turolian/Ruscinian boundary, 16, 615 Venta del Moro fauna, 5ll-513
Turolic faunal unit, Zl Venta del Moro section, 508, 510
Type section, 6 Venta Micena, 349
Verona, 346, 347
Uganda, Z40, 587-589 Veyran, ZZ, Z9
Ulantatar locality, 53Z Vialette, Z4, 36, 1Z5, 1Z6
Untere Landschneckenmergel, Z56 Vialette local fauna, 340
Unterneul, 183, 188 Vienna Basin, Zl4, Zl6
Unterneul clay quarries, 183 Vieux Collonges, 179, 188, 190, 365,
Upper Freshwater Molasse (see also 383, 385, 6Zl
Freshwater Molasse), 178, 179 Viladecavalls, 360, 379, 397
Upper Marine Molasse, 177-180 Villafeliche, 365
Upper Valdarno, 34Z, 344-346 Villafranca d'Asti, 89, 131, 615
Ural Mountains, 410 Villafranchian association, 13Z
U ralian Sea, 548 Villafranchian fauna, 90,347,475-477,
Urungu River, 543 480, 483, 484, 5Z9
Villafranchian faunal expansion, 136
Valdai glaciation, 480 Villafranchian holdover taxa, 349
Valdarno, 131, 13Z, 135, 136 Villafranchian land mammal age, 9, 10,
Valdecebro, 99, 109 91, ll9, 1Z7, 1Z8, 131-136,
Valdemoros, 359 Z74, 305, 346, 356, 547, 615
Valec, Z13, Zl4 Villafranchian sites, 478 (map)
Valerymys-Hispanomys Superzone, 98, Villafranchian stage, Z5
109 Villafranchian-Galerian transition, 347,
Valles de Fuentiduena, 360 356
Valles-Penedes area, 615 Villafranchian-Villanyian land mammal
VaUes-Penedes Basin, 357-360, 365, age, Z4
366, 37 5-379, 381, 383, Villafranchian-Villanyian stage, 36-38
384-387,391-394,396,397, Villafranchic faunal unit, Zl
497, 498 Villalba, 370
Vallesian assemblage, Z30, Z31 Villalba Alta, 35, llO, 399
Vallesian fauna, 57, Zll Villany, Z3Z, 405
Vallesian hipparions, 334 Villany 3, 101, lOZ
Vallesian land mammal age, Z3, 70, 71, vmany 5, 104
90, 94, 96, 108, 11 Z, 166, Zl4, vma.ny 8, 103
Z74, Z96, Z97, 310, 31Z, 3Z8, Villanyian assemblage, Z3Z
330-333, 335, 336, 357, Villanyian fauna, Z14
359-361, 369, 37Z, 375, 383, Villanyian land mammal age, 9, 101,
384, 387, 391, 396,397, 401, 106, 108, llO, 1Z8, 131, 13Z,
40Z, 50Z, 517, 5Zl, 544, 574, Z05, Z06, Z17, 380,405
615, 6ZO, 6Z5 Villanyian stage, Z5
Valles ian localities, 397, 496 (map) Villanyic faunal unit, Zl
Vallesian rodent fauna, 105 Villaroya, Z4, 37, 305
Vallesian stage, 6, 7, Z5, 31, 3Z, 5Z, 53, Villastar, 361
6Zl Vivero de Pinos, 360
Vallesian zone, ZZ7 Voigtsted t, 104
Vallesian zones H and I, 51 V6sendorf (see also Jnzersdorf), Z3, 31,
Vallesian/early Turolian sites, 396 Z30
Vallesian/Turolian boundary, 367, 5Z1 Vrica section, 136
649
Wadi Faregh fauna, 594 Youshashan Formation, 544
Wafangyingzi, 536 Yu County, 547
Wallace's Rule, 418-420, 423, 429, 432 Yunnan Province, 527, 530, 544, 545,
Wangdaifuliang, 545 549, 550
Wasatch Formation, 602, 604 Yushe, 529
Wasatchian land mammal age, 7, 606 Yushe Basin, 532
Wasatchian stage, 7 Yushe County (Shanxi Province), 546
Wattwil, 179 Yushe series, 5Z9
Wattwil fauna, 179
Westbury Cave, 49Z Zagreb, 548
Western Mediterranean, 20 Zalesiaki, Z04
Wet screening, 612 Zamkowa Dolna Cave, Z04, ZOS
Wrrie, 109, 193-195, 202, 204, 205, 232, Zanclean/Piacencian boundary, 20
407, 409 Zhangbei, 536, 537
Wyze 1, 204 Zhangbei County, 536
Wintershof-West, 55, 82, 538 Zhanjiaping, 551
Wisconsin glaciation, 480 Zhanj iaping fauna, 53 7, 551
Wolfersheim, 405, 407, 409 Zhongning County, 545
Wood Committee Report, 606-609 Zhurihe railway station, 544
Woodburne volume, 608 Zirany, Z17
Woodland and forested habitat, 560 Zofingen, 178
Wu Lan Kou (Fugu County), 527 Zonation of Aguilar, 1Z3, 1Z4
Wu-tao-ya-ku, 532 Zonation of Agust1 et al., 1 Z4
Wulungu Formation, 535 Zonation of Cordy, 1Z4
Wiirm glaciation, 480 Zone boundaries, 125
Wurmian glacial fauna, 1Z Zoogeographic region, 549
Wurmian glaciation deposits, 110 ziijar, 381
Wuxiang, 303
Wyoming, 602, 604
Xiacaowan Formation, 68, 538

Xianshuihe, 529
Xianshuihe Formation, 537, 541
Xiaodian Village, 539
Xiaolongtan fauna, 544
Xi~ia, 532, 536, 540, 551
Xiejia fauna, 535
Xiejia Formation, 530, 535, 536, 541
Xin-an (Henan Province), 527
Xining Basin, 530, 535, 541
Xining (Qinghai Province), 529
Xinjian Uygur Autonomous Region, 536,
543
Xinjiang, 535
Xmas Quarry, 294-296
Yassioren, 335, 496-498

Yehucheng Formation, 532
Yellow River, 532
Yeni~ubuk anticline, 64
Yeni-Eskihisar 1 fauna, 23, 31
Yeni-Eskihisar pollen zone, 22, 23
Yenikoy, 64, 66, 68, 69, 516
Yenikoy fauna, 70
Yenikoy section, 517
Yeroham Basin, 240, 594
Yindirte, 538
Yongdong County, 540
Youhe, 547
Youhe fauna, 547
Youhean land mammal age, 274, 304
650
TAXONOMIC INDEX (Mammal Genera)
Aceratherium, 81, 84-87, 140, ZOO, ZOZ, Anomalomys, 85, 86, 96, 98, 108, 179,
Z03, 379, 449, 450, 591 193, 198, Z03, Z11, ZZZ, ZZ7,
Acerorhinus, 544, 545 ZZ8, Z30, 366, 367, 376, 378,
Acinonyx, 89, 131 379, 385, 395-398
Acteocemas, 358, 3 59, 368 Ansomys, 538, 539
Adcrocuta, 86, 87, 334, 379 Antelope, 545
Adelomyarion, 140, 146, 154 Antemus, 573, 756, 580, 58Z-584
Afrosmilus, 591 Anthracobune, Z38
Agriotherium, Z05, ZOO, 546 Anthracotherium, 140
Albanensia, 85, 86 Antilospira, 547
Albanohyus, 85, 86 Aonyx, 89
Albertona, 68, 70 Apeomys, 8Z
Alicornops, 86 Apodemus, 87, 89, 99-106, 109, 110,
Alilepus, 8 7, 545 367, 376, 379-381, 199, 391,
Allocricetus, lOZ-106, 110, Z06, 379 399, 410, 509, 584
Allophaiomys (see also Microtus Aprotodon, 537, 549
(Allopaiomys)), 110, 1Z4, Z06, Arabosminthus, 595
381 Aratomys, 106, 107
Alloptox, 448, 449, 541, 543 Archaeobelodon, Z39, Z40, Z45, 591
Alloscapanus, 84, 85 Archaeomys, 68, 140
Allosorex, Z31 Archidiskodon, 131, 4 75
Alopecocyon, 84, ZOO Arctomeles, ZOO
Amblycoptus, 196 Armantomys, 375, 383, 385, 39Z, 395
Ameniscomys, 8Z, Zl8 Arsinoitherium, Z38
Amphechinus, 81, 84 Arvernoceros, ZOl, Z06
Amphictis, 81-83 Arvicola, 103-105, 107, 110, 1Z4, 349,
Amphicyon, 81, 8Z, 84-86, 161, 449, 381, 407
450, 45Z, 453, 5Z9, 539, 543, Astrohippus, 339
549, 551 Atlantoxerus, 86, 87, 99, 380, 498, 54Z,
Amphicyonopsis, 85 543
Amphilagus, 81-83, 86, 140, 160, Zl8 ·Aureliachoerus, 83, 358, 359, 368
Amphimachairodus, 87 Austroportax, 360
Amphiperatherium, 81, 8Z, 183
Amphitragulus, 81-83, 358, 359 Baranogale, 87, 89, ZOO,Z04, Z05
Anancus, 89, 131, Z31, Z3Z, 475 Baranomys, 99, 100, 10Z, 106, 110, 198,
Anasinopa, 591, 593 Z04, Z05, 405, 407
Anatolomys, 407 Barberahyus, 360
Anchitheriomys, 54Z, 544 Barytherium, Z38
Anchitherium, 51, 53, 83-85, 153, 161, Bathyergoides, 589
168, 173, Z01-Z03, Z39, 435, Bedenomeryx, 140
449, 450, 5Z8, 530, 538, 539, Beliajevina, 449, 450, 45Z
543, 544, 549, 550, 618, 6Zl Bellatona, 543, 544
Ancylotherium, 436, 540 Beremendia, 89, 196
Andegameryx, 8Z, 83, 160, 168, 358, Birgerbohlinia, 87, 360, 36Z, 379
359 Bison, 340, 349
Blackia, 84, 85, 197, Zl8, 410
651
Blancomys, 106, 110, 367, 399, 509 Cormohipparion, 4, Z3, Z68, Z75,
Blarinella, 84, 87, 195 Z90-Z9Z, Z94-Z96, 30Z, 308,
Blarinotdes, 89, 195, 2.31 310, 314, 496
Borissiakia, 537 Coryphodon, 60Z
Borsodia, 94, 96, 97, 99, 101-103, 106, Cremohipparion, Z69, Z75, Z77, Z78,
110, ZOS, Z3Z, 407, 408 Z9Z, Z97, 307-314
Brachyerix, 535 Cricetodon, 67, 68, 70, 72., 84, 85, 183,
Brachyodus, 160, 161, 168, 173, Z39, 184, 188, 190, zzz, Z37,
Z55, 358, 368, 371, 435, 436, 365-367' 377' 378, 385-387'
591 391, 394, 395, 397, 498, 54Z
Brachypotherium, 8Z, 83, 84, ZOO, ZOZ, Cricetulodon, 86, 360, 367, 376, 378,
Z03, 449, 450, 535, 536, 539, 379, 497' 498
548, 591, 593 Cricetulus, 7Z
Brachyrhizomys, 530, 545, 546, 580 Cricetus, 68, 72., 87, 99, 104-106, 110,
Bransatoglis, 66, 70, 81-83, Z18, ZZZ, Z06, 367' 380, 381, 387' 509
366, 377' 378, 383, 384, 393, Crocuta, 347, 4Z5
394, 396 Croizetoceros, 89, Z01, ZOS
Broiliana, 83, 160, 173 Crouzelia, 84
Bubalus, 1Z6 Crusafontina, 86
Bunolistriodon, 83, 173, Z58, 359 Cseria, 406, 409
Byzantinia, 68, 106, 109, 497, 498 Cynelos, 8Z, 160
Cadurcotherium, 140 Dakkamys, 576

Cainotherium, 81-83, 140, 183, 2.54 Dama, 133, 135
Calomyscus, 106, 367, 387, 401 Decennatherium, 87, 334, 360
Camelus, 436 Deinosorex, Z2.7
Canis, 87, 1Z4, 133-136, 346,448,480, Deinotherium, 83, 84, 86, 166, 173, 181,
491 190, Z11, Z37, Z53, Z54,
Canthumeryx, Z40, 591, 593 Z56-Z59, Z61, 449, 450, 549
Capreolus, 86 Deinsdorfia, 195
Caprotragoides, 359, 360, 368, 371, 449, Democricetodon, 5Z, 53, 55, 57, 66, 67,
451, 596 7Q-73, 83-86, 96, 98, 107, 108,
Carposorex, 8Z 183, 190, 193, 198, zoz, Z03,
Castillomys, 70, 101, 367, 379, 381, ZZZ, ZZ4, ZZS, ZZ7, 2.30, Z37,
391, 400 365, 367' 369, 377' 386, 392.,
Castor, 89 394, 448, 449, 537, 538, 543,
Celadensia, 99, 100, 367, 407 549, 551, 576, 577, 583
Cephalogale, 81, 140 Dendromus, 106
Cercopithecus, 419, 4ZZ Deperetomys, 70, 7Z, 85, 105, 365, 385
Cervalces, 346, 347 Desmanella, 85, 87, 54Z
Cervavitus, 547 Desmana, 19 5
Cervus, 89, 135, Z01, ZOS, 349, 480 Desmanodon, 44 7-449
Chalicomys, 98, 198, Z03 Desmatolagus, 536
Chalicotherium, 84, 86, 173, Z01, ZOZ, Diaceratherium, 81, 83, 140, 160, 536
379, 449, 539, 541, 543, 591 Diatomys, 538, 539
Chardinomys, 547 Dibolia, 87
Chasmaporthetes, 89, 131, 334, 546 Diceratherium, 81
Chilotherium, Z60, 540, 541, 545, 549 Dicerorhinus, 81, 86, 87, 89, ZOO, Z05,
Chleuastochoerus, 544 379, 540, 541, 591
Choerolophodon, 2.37, 2.39, Z4Z, Z47-Z49 Diceros, 550
Circamustela, 86 Dicoryphochoerus, 544
Citellus, 104, 105 Dicrocerus, 84, 173, 543, 544
Clapasorex, 81 Dicrostonyx, 104, 105, 107, 110
Clethrionomys, 10Z-106, 110, Z3Z Dimylechinus, 81, 535
Collimys, 96, 98, 105, 108 Dimylus, 81
Collongomys, 365 Dinocrocuta, 544, 545
Condylura, 195 Dinofelis, 135
Conohyus, 85, 166, Z01, Z03, 368, 449, Dinohippus, 339
451, 563 Dinosorex, 84-86, ZZ7
Copemys, 386 Dionysophithecus, 538
Cordylodon, 8Z Dipoides, 86, 87, 99
652
Dolomys, 101, 102., 106, 109, 110, 198, Fortunictis, 87
2.04, 2.05, 407
Dorcabune, 546 Gaindatherium, 173
Dorcatherium, 83-87, 173, Z01, Z03, Gaindatherium (Iberotherium), 2.56, Z58
332., 334, 358, 361, 368, 371, Galemys, 195
449, 451, 538, 551, 591, 592. Galerix, 83-87, ZZ7
Dremotherium, 81, 82., 140 Gallogoral, 89
Dryomimus, 10Z, 103 Gazella, 87, 89,475, 545, 547
Dryopithecus, 86, 376, 447, 530, 549 Gazellospira, 89
Dzungariotherium, 536, 537 Gelocus, 591
Germanomys, 101, 102., 106, 109, 198,
Ebromys, 4 71 2.04, Z31, 405, 407, 408, 547
Elephas, 439, 547, 550, 551, 563, 604 Giraffa, 563
Eliomys, 86, 87, 98,99, 105, 2.Z7, Z30, Giraffokeryx, 449, 451
2.31,366,380, 383,384,399, Gliravus, 140, 154
409, 410, 508 Glirudinus, 66, 70, 81-83, 85, 86, 96,
Enhydrictis, 89 199, 2.18, zzz, 377, 383, 393,
Enhydriodon, 87 394
Entelodon, 548 Glirulus, 89, lOZ, 199, ZZZ, ZZ7, Z30,
Eolagurus, 105, 107, 110 405, 410
Eomuscardinus, 84-86, 2.2.7, 366, 383, Glis, lOZ, 105, 199, ZZZ, ZZ7, Z30, 376,
394, 396 383, 384, 396, 409, 410
Eomyops, 85, 86, ZZO, ZZZ, Z2.7, Z31, Glyptotherium, 604
366,369,376,378,396,398 Gobicyon, 540, 543, 549
Eomys, Z18 Gomphotherium, 83-85, 181, Z01-Z03,
Eostylocerus, 87, 360, 361, 379 Zll, ZZZ, Z37-Z4Z, 2.45, 2.48,
Eotragus, 84, 85, Z37, 2.40, 359, 360, Z54, Z56, Z59, Z61, 449, 450,
368, 371, 540 453, 537' 538, 541, 548-550,
Eozapus, 98, 99 591
Eozygodon, Z41, 2.42., 2.45, 591, 59Z Graphiurops, ZZ7, Z30, Z31
Epimeriones, 96, 98, 100, 105, 198, Z04,
367, 376, 390, 400 Haplocyon, 81, 8Z, 140
Episoriculus, 89, 196 Haplocyonoides, 8Z, 160
Eptesicus, 84. 196 Haplocyonopsis, 81
Equus, 9, 89, 131, 305, 339-353, 435, Hecubides, 591
436, 475, 480, 550, 551, 563 Hemicyon, 83-85, 449, 539, 540, 543,
Erinaceus, 195, 544 591
Estramomys, lOZ, 103, 197, Z04, Z05, Hemimastodon, Z37-Z39, 2.41, Z4Z, 2.44,
366, 399 Z49
Eucladoceros, 133, 135, 2.05, 34Z, 346 Herpestides, 81, 8Z, 591
Eucricetodon, 5Z, 53, 55, 57, 67, 68, 70, Heterictis, ZOO
n, 81, 8Z, 140, 146, 147, 154, Heteromyoxus, 8Z, Z18
160, 178, 365, 39Z, 535, 536, Heteroprox, 84, 85, 357, 360
548 Heterosorex, 8Z
Euctenoceros, 89 Heteroxerus, 81-84, 86, 87, 2.18
Eumyarion, 66, 70, 7Z, 83-86, 96, 105, Hexaprotodon, 1Z6, 563
108, 179, 183, 186, 190, 193, Hipparion (see also Cormohipparion and
198,Z03, Z11, ZZZ,ZZ3, ZZ7, Hippotherium), 4, 9, 10, 19,
Z30, 365, 367, 369, 377, 379, Z3, 51, 53, 54, 57, 86, 87, 90,
392., 395, 396 96, 166, 173, ZZ7, Z48, Z54,
Euprox, 85-87, 193, ZOl-2.03, 359, 360, Z68, 2.75, Z78, Z9Q-2.93,
368, 449, 451, 543 2.95-Z99, 301-310, 31Z, 314,
Eurolagus, 84, 85, 197, ZOZ 315, 32.1, 3Z8-330, 33Z, 333,
Eutamias, 538 336, 360, 378, 379, 383, 386,
Exallerix, 535 395, 436, 496, 497, 500, 502.,
503, 52.7-530, 53Z, 544-547,
Fahlbuschia, 53, 105, 360, 365, 367, 550,551,576,597,618
369, 377, 378, 3.94 Hipparion s.s., Z76, Z98, 30Q-30Z, 307,
Felis, 89, ZOO, Z05 311-315
Felsinotherium, 89 Hippopotamus, 361, 369, 371, 4Z5, 436
Forsythia, 85 Hipposideros, 83
653
Hippotherium, Z3, Z15 Leakitherium, 591
Hippotigris, 344, 348 Legetetia, 591
Hispanodorcas, 87, 361 Lemmus, 94, 97, 10Z-107, 110, Z06
Hispanomeryx, 359, 360, 368 Leptarctos, 543
Hispanomys, 86, 98, 99, 105-109, 367, Leptobos, 89, 131, 133, 475
378-380, 385, 394, 395, 397, Leptodontomys, 197, Z04, 54Z
398, 498 Leptop1esictis, 85, 591
Hispanotherium, 173, Z56, Z58, Z60, Leptotataromys, 53Z, 536
Z61, 359, 530, 541, 54Z Lepus, 436
Homoiodorcas, 369, 596 Leukaristomys, 407
Homotherium, 89, 131 Libycochoerus, 591, 593
Hyaena, 133, 33Z Ligerimys, 5Z, 53, 8Z, 83, 153, 154, 178,
Hyaenaelurus, 549, 589 179, Z18, Z19, ZZZ, 377, 39Z,
Hyaenodon, 140,537,591 393,471
Hydracoides, 86 Listriodon, 84-86, 359, 360, 368, 409,
Hydromops, 81 436, 450, 451, 5Z9, 530, 540,
Hyoboops, 591 54Z, 543, 563, 593
Hyotherium, 81-83, 86, 359, 360, 368, Lophiba1uchia, 69, 70
544 Lophiodon, 613
Hypolagus, 89, 197, Z05, Z31 Lophiomeryx, 536
Hypsodontus, 451 Lophocricetus, 407
Hyracotherium, 60Z Lufengpithecus, 546
Hystrix, 90, ZOO, Z04, 436, 567, 576 Luoga1e, 591
Lutra, 546
Ictitherium, 334, 544, 546 Lycyaena, 87
Indarctos, 86, 87, 546 Lynx, 475
Ischymomys, 106, 107, 109
Ischyrictis, 84-86 Macaca, 89, 369, 371, 4Z5, 540
Isoptychus, 613 Machairodus, 86, 87, 543, 546
Issiodoromys, 68, 140 Macrotherium, 165, 540, 541
Mafia, 195
Kansupithecus, 538 Mammut, Z01, Z31, Z3Z, 341, 546, 547
Karnimata, 367, 391, 58Z, 584 Mammuthus, 89
Kelba, 589, 591 Marcetia, 86
Kenyalutra, 591 Martes, 83-86, ZOO, Z05, 543
Kenyapithecus, 44 7 Megaceros, 349
Kenyasus, 591 Megacricetodon, 53, 67, 70, 7Z, 83-86,
Keramidomys, 84-86, 197, ZZO, ZZZ, 105, 108, 179, 183, 185,
ZZ7, Z31, 366, 376, 394-396, 187-190, zzz, ZZ6, ZZ7, Z30,
398, 399, 54Z Z37, 365, 367, 369, 377-379,
Keridomys, Z04 383, 385, 386, 39Z, 394-396,
Kichechia, 591 448, 449, 497, 498, 537, 538,
Kipsigicerus, 369 541, 543, 549, 551, 576, 583,
Kislangia, 379, 381 6Z1 .
Kowalskia, n, 86, 87, 98-101, 105, 108, Megaderma, 8Z, 87, ZZ7
110, 198, Z04, ZZ4, ZZ5, ZZ7, Megalovis, 1Z4, 136, 346
Z30, Z31, 367, 376, 380, 387, Megamphicyon, 8Z, 83
397, 398, 409, 410 . Megantereon, 89, 131
Kubanochoerus, Z48, 368, 540-543, 549, Megapedetes, Z48, 589
551 Megistotherium, 591
Kubanotragus, 449, 451, 453 Meinia, 539
Meles, 89
Lagomeryx, 83, 160, 168, 173, 358, 359, Melissiodon, 5Z, 8Z, 83, 140, 179, Z18,
538, 539, 543, 544, 551 ZZ1, ZZZ, 39Z
Lagopsis, 83-86, 145, 154, 160 Melodon, 543
Lagurodon, 10Z-104, 106, 110 Merychippus, Z68, Z90, Z91, 30Z
Lagurus, 105, 107, 110 Mesaceratherium, 81, 83
Lanthanotherium, 83-86, ZZ7 Mesembriacerus, 331, 334
Laphyctis, 83 Mesochoerus, 439
Lartetomys, ZZZ, 365, 385 Mesocricetus, 68, 7Z
Lartetotherium, 83-85, 541 Mesomephitis, 86
654
Metacordylodon, 85, 86, 195, ZOZ Myorycteropus, 591
Metailurus, 87, 543, 546 Myotis, 81-84, 196
Metapterodon, 591
Metasayimys, 594 Nannippus, Z75
Metaschizotherium, 540 Necromanis, 82., 83
Metexallerix, 53Z, 535 Neocometes, 85, 105, 193, Z11, ZZ.Z,
Microbunodon, 140 ZZ7' ZZ9, 369
Microdyromys, 81, 83-85, Z18, ZZZ, 384, Neocricetodon, 387
393, 538, 54Z Neodon, 104, 105
Micromeryx, 84-86, 361, 368, 449, 451, Neohipparion, Z75, 30Z, 305
541, 543 Neomysorex, 196
Micromys, 99, 100, 106, 199, 379, 409, Neurotrichus, 195
410, 584 Ningxiatherium, 545
Microstonyx, 87, 360, 361, 379 Nisidorcas, 332., 334, 369
Microtia, llZ Notiocradohipparion, Z91, Z92.
Microtocricetus, 96, 98, 105, 107, 108, N otochoerus, 439
386, 387, 397 Numidotherium, Z38
Microtodon, 106, 198, Z04, 407 Nyctereutes, 89, 134, ZOO, 475, 546,
Microtoscoptes, 106, 107, 109, 407 547, 550
Microtus, 103-105, 110, 2.06 Nyctinomus, 81, 85
Microtus (Allophaiomys), 94, 101, 107,
2.32. Occitanomys, 70, 87, 96, 98-100, 105,
Mimomys, 89, 90, 100, 101, 106, 107, 109, 11Z, 367' 379-381, 391,
109, 110, 1Z4, 131, 13Z, 198, 398, 399, 508
199, 2.04-Z06, Z31, Z3Z, 381, Ochotona, 197, ZOS, 547
405, 406, 408-411, 547 Ochotonoides, 54 7
Mimomys (Cseria), 94, 101-103 Oioceros, 369, 535, 536, 540-543, 595
Mimomys (Kislangia), 10Z Oligosorex, 81, 84
Miniopterus, 196, 2.04 Olobulukia, 544
Miodyromys, 8Z, 84, 85, 2.18, ZZZ, 377, Orientalomys, 100, 101, 547
383, 395 Oriomeryx, 8Z
Mioechinus, 84 Orycteropus, 89, 2.48, 449, 451, 550,
Miomachairodils, 544, 545 591, 592.
Miomegaderma, 84, 85 Oryctolagus, 89
Miomephitis, 83 Ouranopithecus, 334
Mionictis, 84, 543 Ouzocerus, 334
Miopetaurista, 70, 8Z-86, 197, Z18, 377
Mioprionodon, 591 Pachycrocuta (see also Pliohyaena), 89
Miorhynchocyon, 589 Pachyhyrax, 591
Miosorex, 83, 84, 86 Pachytragus, 451, 596
Miotragocerus, 86, 332., 359, 360, 368, Paenelimnoecus, 82., 84, 87, 196
369 Palaeochoerus, 82., 140
Mirabella, 66, 70 Palaeogale, 81-83
Moeritherium, 2.38 Palaeomastodon, 2.38
Monosaulax, 536, 54Z Palaeomeryx, 83-85, Z01, ZOZ, 357, 358,
Montalbanensis, 70 368,449,451,538-542.
Moropus, 81 Palaeomys, 86
Mus, 107, 110, 576, 577 Palaeonycteris, 81
Muscardinus, 85-87,89,98,99, 105, Palaeoreas, 334
199,ZZ7, 2.30, 366,367,369, Palaeoryx, 369
376-380, 383, 384, 394, 396, Palaeosciurus, 70, 81-83, 197
410 Palaeotapirus, 81, 539
Mustela, ZOO, Z05 Palaeotragus, 360, 368, 538, 541-544,
Mygalea, 84 547
Myocricetodon, 106, 367, 371, 387, 401, Pannonicola, 106
576 Panthera, 89, 1Z6, 133
Myoglis, 84-86, 96, 98, Z18, ZZZ, ZZ7, Paracamelus, 361, 546, 550
Z30, 366, 369, 378, 383, Paracervulus, 546
394-396 Parachleuastochoerus, 86, 360
Myomimus, 71, 87, 98,366, 369,379, Paracitellus, 82.
381,383,396,399,498 Paraentelodon, 537, 548
655
Paraethomys, 100, 101, 106, 109, 110, Pliomys, 101, 104, 105, 107, 109, 110,
367' 376, 380, 381, 391, 399, 379
509, 577, 582, 584 Pliomys (Propliomys), 106
Paraglirulus, 85, 86, 227, 230, 231, 366, Pliopentalagus, 231
369,378,383,384,394,396, Pliopetaurista, 102, 197,231,409,410
398 Pliopetes, 102, 197
Paraglis, 84, 85, 227 Pliopithecus, 83, 84, 165, 173, 183, 196,
Parahippus, 268, 290, 291 202, 203, 540
Parailurus, 89, 231, 232 Plioselevinia, 199
Paralactaga, 542 Pliospalax, 7Z, 96, 105, 109, 448, 449
Paralutra, 85, 86 Plioviverrops, 87, 334, 379, 550
Paramachairodus, 87 Plithocyon, 84,160,449
Paranomalurus, 589 Postpalerinaceus, 86
Paranourosorex, 196 Potamotherium, 81
Parapelomys, 584 Potwarmus, 576
Parapetaurista, 538 Praearmantomys, 375, 377
Paraphiomys, 589 Praemegoceros, 349
Parapliohyrax, 451 Pristiphoca, 2.2.7
Parapodemus, 68, 70, 72, 87, 90, 98, 99, Proailurus, 81, 82
105, 109, 199, 227, 231, 360, Proboscidipparion, 269, 275, 303, 304,
361, 367, 379, 380, 391, 398, 306, 307, 312, 314, 546, 547
508, 509, 511, 582, 584 Procamptoceras, 124, 136
Parasminthus, 68, 535 Procapreolus, 201, 205, 227, 361
Parasorex, 86 Procervulus, 83, 358
Paratalpa, 81 Proconsul, 589
Paratapirus, 161 Prodeinotherium, 86, 238, 240, 241,
Pelomys, 106 245, 591-593
Pelycodus, 602 Progenetta, 83
Percrocuta, 449, 450, 530, 540, 543 Progiraffa, 368, 371
Peridyromys, 81-83, 140, 218, 222, 377, Progonomys, 68, 70, 7Z, 86, 98, 105,
381,383, 39~ 393,471 108, 331, 334, 360, 367 376,
Petenyia, 89, 195 379, 390, 396, 497' 498, 502,
Petenyiella, 380 574, 576, 580, 582-584
Phenacolophus, 238 Prohyrax, 591
Phiomia, 238, 242, 244 Prokanisamys, 575
Phoberocyon, 83 Prolagurus, 104, 107, 110
Phyllotillon, 81, 85, 537, 548, 549 Prolagus, 82.-87, 89, 145, 154, 160, 179,
Piezodus, 81, 140 379, 448, 449
Pitymys, 104, 105 Promephitis, 86
Platodontopithecus, 538 Prometheomys, 2.04
Platybelodon, 248, 528, 532, 540, 541, Promimomys, 68, 70, 7Z, 99, 106, 109,
543, 545, 549, 551, 591 110
Plecotus, 196 Proochotona, 498
Plesiaceratherium, 83, 529, 539, 540, Propalaeoryx, 591
551 Propliomys, 101, 199, 204, 205, 407, 408
Plesiaddax, 369, 549 Propotamochoerus, 544
Plesictis, 81-83, 140 Proputorius, 84, 85, 449, 546
Plesiodimylus, 82-86, 227 Prosantorhinus, 83
Plesiodipus, 541, 542 Proscapanus, 82-84
Plesiogale, 81-83 Prosiphneus, 544, 547
Plesiogulo, 334, 449, 546 Prosomys, 198, 204
Plesiosminthus, 81, 140, 146, 154, 535, Prospalax, 98, 102-105, 109, 110, 199,
538, 539, 548 408
Plesiosorex, 85 Prostrepsiceros, 33 2, 334
Plesippus, 340 Protaceratherium, 81, 83, 160
Plesispermophilus, 82., 2.18 Protalactaga, 541, 542, 544
Plioctomys, 407 Protanancus, 245
Pliohyaena, 133, 546 Protapirus, 140
Pliohyaena (= Pachycrocuta), 131 Protatera, 106,367,371,381,387-390,
Pliohyrax, 86, 451 401, 576
Protictitherium, 86, 449, 450, 538
656
Protoryx, 369 Selenoportax, 563
Protozapus, 227, 231 Semigenetta, 83-86, 160, 173, 538
Protrage1aphus, 332 Serridentinus, 543
Protragocerus, 360 Shaanxispira, 545
Protursus, 86 Shamalina, 595
Pseudaelurus, 83-85, 200, 203, 449, 538 Shuanggouia, 53 8
Pseudarctos, 83-85, 543 Sic~ta, 104
Pseudocricetodon, 67, 68, 70, 72, 82, Simamphicyon, 86
140 Sinoadap~, 545
Pseudocyon, 81-84 Sinolagomys, 535, 548
Pseudocyonops~, 81, 140 Sinomioceros, 543
Pseudodryomys, 55, 81-83, 140, 222, Sivalhippus, 268, Z75, Z82, 303-307
227,377,381, 383,39~393, Sivameryx, 436
449, 471, 472 Sivaonyx, 87, 546
Pseudofahlbuschia, 53 Sivapithecus, 443, 44 7, 449, 544, 546,
Pseudogalerix, 84 549, 567
Pseudohipparion, 275 Sivoreas, 369, 371
Pseudo1tinomys, 68 Sminthozapus, 10Z, ZOO, Z04, Z05, 367,
Pseudomeriones, 71, 72, 100, 108, 230, 376, 400, 410
367' 390, 401 Soergelia, 349
Pseudotheridomys, 81, 82, 140, 150, Sorex, 8Z, 89, 195
152, 178, 218, 219, 222, 471 Soricella, 8Z
Pterodon, 591 Spalax, 104, 107
Ptychoprolagus, 218 Spanocricetodon, 66, 67, 70-72, 386,
39Z, 538, 539, 575
Quercomys, 471 Spermophilinus, 83-87, 99, 107, Z18,
Qurliqnoria, 544, 545 ZZ7, 54Z
Spermophilus, Z06
Rahonapithecus, 86 Stachomys, 101, 10Z, 106, 109, 198,
Ramapithecus, 447, 546, 549 204, Z05, 408, 411
Rangifer, 482 Stenailurus, 379
Rattus, 107, 110 Stegodon, 546, 563
Ratufa, 82 Steneofiber, 81-84, 86, 160, 173, 198,
Renzimys, 53 536
Rhagapodemus, 100, 106, 110, 112, 199, Stenogale, 81
231, 379, 391, 409, 410 Stenoplesict~, 84, 591
Rhino1ophus, 81, 82, 196, 204 Stephanocemas, Z39, 360, 538, 541, 543,
Rhizomys, 449 544
Rhizospalax, 140, 154 Stephanochoerus, 543
Rhodanomys, 81, 140, 146, 147, 149, Stephanomys, 87, 99-101, 109, 110, 11Z,
154, 471, 47Z 361, 367' 379-381, 391, 399,
Ritteneria, 52, 149, 154, 47Z 509
Rotundomys, 105, 107, 108, 367, 379, Strogulognathus, 84
386, 387' 396-398 Stromeriella, 83, 160, 173
Ruscinomys, 87, 94, 99, 100, 106, 109, Stylohipparion, Z75, 306, 307
110, 112, 367, 381, 385, 398, Sulimskia, 19 5
399, 509 Sus, 89, 133, 341, 349, 547
Synaptomys, 94, 97, 10Z, 106, 110, 199,
Sabadellictis, 86 205
Samotherium, 544
Sansanosmilus, 84, 85, 540, 541 Tachyoryctoides, 535, 537, 548
Sayimys, 68, 71, 538, 549, 575 Tadarida, 85
Scapanulus, 19 5 Talpa, 8Z-87, 89
Scaptonyx, 85, 195, 227 Tamias, 71, 197, 409, 410
Schizochoerus, 360, 379, 397 Tapirus, 83, 86, 89, ZOO, Z03, ZZ7, Z31,
Schizogalerix, 380, 447-449, 498 Z3Z, 341, 476, 544
Schlossericyon, 84 Tataromys, 67, 53Z, 53 5, 536, 548
Sciuropterus, 85 Tatera, 389, 390
Sciurus, 84, 197, 542 Taucanamo, 84,Z01, Z03,449,451
Scotophilus, 85 Taxodon, 84
Selenolophodon, 541, 542
657
Tempestia, 86, 98, 366, 369, 375, 395, Valerymys, 87, 96, 98, 99, 105, 106,
396 109,367,37 9,380,391,3 98
Teratodon, 589, 591 Vasseuromys , 66, 70, 81, 8Z, 154, 471
Teruelia, 358, 368 Villanyia, 94, 96, 97, 99, 101-103, 110,
Tesselodon, 543 199, Z05, Z3Z
Tetralophod on, 86, 87, Z01, 544, 545 Viverra, 84, 89
Thalassictis, 87 Vulpes, 89, 134, 448, 490
Titanomys, 81, 145, 154, 536, 537
Tragocerus, 87 Walangania, 591
Tragoportax , 87,334,360- 36Z, 369,
371, 379 Xenochoerus , 591, 593
Triceromery x, 357, 359 Xenocyon, 135
Trilophomys , 94, 98-100, 101, 106, 110, Xenohyus, 8Z, 160, 358
198, Z04, Z05, Z31, 381, 405,
407, 408 Youngofiber , 538
Trocharion, 85, 86 Ysengrinia, 81, 8Z
Trochictis, 86, 165, 449 Yunnanother ium, 546
Trochotheriu m, 85
Trogontheriu m, 84-86, Z05 Zelceina, 195
Tsaganomys , 53Z, 548 Zygolophodo n, 89, 131, Z37-Z39, Z48,
Tsaidamorth erium, 544 340, 543, 547
Tungurictis, 543
Turcocerus, 543
Turkomys, 67, 448, 449, 497
Ungaromys, 101-103, Z04, Z3Z, 405,

407, 547
Urmiatheriu m, 550
Ursavus, 83, 85, 86, ZOO, 449, 539, 546,
551
Ursus, 89, ZOO, 341, 546
658

1989 European Neogene Mammal Chronology

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1989 European Neogene Mammal Chronology

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European Neogene

The series is published by an international board of publishers in conjunction with the

A Life Sciences Plenum Publishing Corporation

c Mathematical Kluwer Academic Publishers

F Computer and Systems Sciences Springer-Verlag

Recent Volumes in this Series

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Volume 176-Processing of Sensory Information in the

Volume 177-Prostanoids and Drugs

Volume 178- The Enzyme Catalysis Process: Energetics, Mechanism,

Volume 179-lmmunological Adjuvants and Vaccines

Volume 180-European Neogene Mammal Chronology

Series A: Life Sciences

Springer Science+ Business Media, LLC

Llbrary of Congrass Cataloglng-ln-Publlcat ton Data

European Naogane •ammal chronology 1 edited by Everett H. Lindsay,

© 1989 Springer Science+Business Media New York

It is fitting and proper that a volume on European Neogene mammal chronology

The rich record of fossil mammals in Europe has contributed substantially to

An international workshop was held at Schloss Reisensburg in Germany on May

Contributions to this volume reflect the diversity of topics and viewpoints

ASPECTS OF EUROPEAN MAMMAL CHRONOLOGY

European Neogene Marine/Continental Chronologie Correlations • • • • • • • • • • • • • 15

A Biochronologic Subdivision of the European Paleogene Based on

The Ramblian and Aragonian: Limits, Subdivision, Geographical

New Neogene Rodent Assemblages from Anatolia (Turkey) 61

Muroid Rodent Biochronology of the Neogene and Quaternary • • • • • • • • • • • • • • • • 91

Biozones or Mammal Units? Methods and Limits in Biochronology • • • • • • • • • • • • 119

Synthesis on the "Aquitanian" Lagomorph and Rodent Faunas of the

The Faunas and Stratigraphical Subdivisions of the Orleanian

The Faunal Succession in the Bavarian Molasse Reconsidered-

Stratigraphy of Neogene Mammals of Poland 193

The Neogene VP Sites of Czechoslovakia: A Contribution

FAUNAL DATUM PAPERS

The "Proboscidean Datum Event:" How Many Proboscideans

The Proboscideans Data, Age, and Paleogeography: Evidence

Patterns of Old World Hipparionine Evolutionary Diversification

The Hipparions of the Lower Axios Valley {Macedonia, Greece).

The Genus Eguus in Europe • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 33 9

Bioevents and Mammal Successions in the Spanish Miocene • • • • • • • • • • • • • • • • • • 357

The Miocene Rodent Succession in Eastern Spain: A

Gundersheim-Findling, a Ruscinian Rodent Fauna of Asian

Dynamics of Old World Biogeographic Realms during the

Miocene Paleoecology of Pasalar, Turkey • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 443

Relations Between Paleoclimatology and Plio-Pleistocene

Small Mammal Taphonomy 487

Hipparion Datum and its Chronologie Evidence in the

The Magnetic Stratigraphy of the Late Miocene Sediments

Preliminary Magnetostratigraphic Results of Some Neogene

SEQUENCES OUTSIDE EUROPE

The Chinese Neogene Mammalian Biochronology- Its Correlation

Key Biostratigraphic Events in the Siwalik Sequence •• , •• , ••• , , , •• , • , • • • • • • • 557

Quo Vadis, Antemus? The Siwalik Muroid Record 573

Development and Application of Land Mammal Ages in

The Past, the Present, and the Future 625

Subject Index • • . . . . . . . . . . . . . • . . . . . • . . . . . . . . . . . . . . . . . . • • . . • . . . . . . . . . . . • 631

Taxonomic Index (Mammal Genera) . •• . . . •. . . • . . •. •. . . •. . . . . • . • . . . • . . •. . . 651

Vertebrate paleontology represents a blend of the biologic and geologic

The communication problem is not unique to vertebrate paleontologists, nor is it

In this contribution I review some of the basic strategies used by vertebrate

European Neogene Mammal Chronology

Stratigraphic superposition is the fundamental strategy upon which the Geologic

Fig. 1. Three ways of approaching the

Superposition is virtually a requirement for all biostratigraphic reference